Safety first: terrestrial predators drive selection of highly specific nesting sites in colonial‐breeding birds

Nesting is a critical yet hazardous life stage for many birds. For colonial‐breeding birds, the conspicuousness of the colony to predators suggests immense pressure to select optimal colonial nesting sites. But what drives selection of those sites? As with solitary nesting birds, reducing access by predators may be the single most important factor. If so, knowledge of the predators involved and the attributes of different potential colony sites can allow us to predict the features that make a site especially safe. We examined the attributes of trees used by breeding colonies of metallic starlings Aplonis metallica in tropical Australia, and experimentally tested if those attributes prevented nest access by predatory snakes. Our surveys confirmed that tree choice by starling nesting colonies is highly non‐random, with all colonies located in tall trees in rainforest clearings, with no low branches and smooth bark. Experimental tests demonstrated that the climbing ability of predatory snakes depends upon bark rugosity, and that colony access by snakes depends on tree attributes such as bark rugosity and canopy connectivity. Our study confirms that colonial‐nesting starlings select colony sites that provide a safe refuge from predation. Intense predation pressure may have driven the evolution of stringent breeding habitat criteria in many other species of colonial‐breeding birds.     

Gap‐crossing decisions of forest birds in a fragmented landscape

Habitat loss and fragmentation are recognized as primary drivers of biodiversity loss worldwide. To understand the functional effects of habitat fragmentation on bird populations, data on movement across gaps in habitat cover are necessary, although rarely available. In this study, we used call playback to simulate a conspecific territorial intruder to entice birds to move through the landscape in a predictable and directional manner. We then quantified the probability of movement in continuous forest and across cleared gaps for two forest‐dependent species, the grey shrike‐thrush (Colluricincla harmonica) and the white‐throated treecreeper (Cormobates leucophaeus). Fifty‐four playback trials were conducted for each species across distances ranging from 25 to 480 m in continuous forest and 15–260 m across gaps in a forest‐agricultural landscape in southern Victoria, Australia. The probability of movement was significantly reduced by gaps in forest cover for both species. Shrike‐thrushes were six times more likely to move 170 m in continuous forest than to cross 170‐m gaps. The mean probability that treecreepers would cross any gap at all was less than 0.5, and they were three times less likely to move 50 m across a gap than through continuous forest. Both species displayed non‐linear responses to increasing gap distance: we identified a gap‐tolerance threshold of 85 m for the shrike‐thrush and 65 m for the treecreeper beyond which individuals were most unlikely to cross. The presence of scattered paddock trees increased functional connectivity for the shrike‐thrush, with individuals crossing up to 260 m when scattered trees were present. We conclude that gaps in habitat cover are barriers to movement, and that characteristics of the intervening matrix influence landscape permeability.     

landscapemetrics: an open‐source R tool to calculate landscape metrics

Quantifying landscape characteristics and linking them to ecological processes is one of the central goals of landscape ecology. Landscape metrics are a widely used tool for the analysis of patch‐based, discrete land‐cover classes. Existing software to calculate landscape metrics has several constraints, such as being limited to a single platform, not being open‐source or involving a complicated integration into large workflows. We present landscapemetrics, an open‐source R package that overcomes many constraints of existing landscape metric software. The package includes an extensive collection of commonly used landscape metrics in a tidy workflow. To facilitate the integration into large workflows, landscapemetrics is based on a well‐established spatial framework in R. This allows pre‐processing of land‐cover maps or further statistical analysis without importing and exporting the data from and to different software environments. Additionally, the package provides many utility functions to visualize, extract, and sample landscape metrics. Lastly, we provide building‐blocks to motivate the development and integration of new metrics in the future. We demonstrate the usage and advantages of landscapemetrics by analysing the influence of different sampling schemes on the estimation of landscape metrics. In so doing, we demonstrate the many advantages of the package, especially its easy integration into large workflows. These new developments should help with the integration of landscape analysis in ecological research, given that ecologists are increasingly using R for the statistical analysis, modelling and visualization of spatial data.     

Recent population declines in Afro‐Palaearctic migratory birds: the influence of breeding and non‐breeding seasons

Aim

Recent, rapid population declines in many Afro‐Palaearctic migratory bird species have focussed attention on changing conditions within Africa. However, processes influencing population change can operate throughout the annual cycle and throughout migratory ranges. Here, we explore the evidence for impacts of breeding and non‐breeding conditions on population trends of British breeding birds of varying migratory status and wintering ecology.

Location

Great Britain (England & Scotland).

Methods

Within‐ and between‐species variation in population trends is quantified for 46 bird species with differing migration strategies.

Results

Between 1994 and 2007, rates of population change in Scotland and England differed significantly for 19 resident and 15 long‐distance migrant species, but were similar for 12 short‐distance migrant species. Of the six long‐distance migrant species that winter in the arid zone of Africa, five are increasing in abundance throughout Britain. In contrast, the seven species wintering in the humid zone of Africa are all declining in England, but five of these are increasing in Scotland. Consequently, populations of both arid and humid zone species are increasing significantly faster in Scotland than England, and only the English breeding populations of species wintering in the humid zone are declining.

Main conclusions

Population declines in long‐distance migrants, especially those wintering in the humid zone, but not residents or short‐distance migrants suggest an influence of non‐breeding season conditions on population trends. However, the consistently less favourable population trends in England than Scotland of long‐distance migrant and resident species strongly suggest that variation in the quality of breeding grounds is influencing recent population changes. The declines in humid zone species in England, but not Scotland, may result from poorer breeding conditions in England exacerbating the impacts of non‐breeding conditions or the costs associated with a longer migration, while better conditions in Scotland may be buffering these impacts.

     

Determinants of species‐area relationships for marsh‐nesting birds

ABSTRACT To clarify the underlying causes of the species‐area relationship in marsh‐nesting birds, I studied eight freshwater tidal marshes of the Connecticut River that differed in area, degree of isolation, mudflat cover, water cover, tidal regime, and extent of individual plant communities. I measured these habitat variables on aerial infrared photos, and surveyed bird populations by mapping the distribution of all birds in marshes under 5 ha in area and establishing 50‐m radius plots in marshes over 5 ha. From surveys, I determined species richness, population densities, and total populations. Analysis revealed a positive relationship between species richness and area, but no correlation between area and habitat heterogeneity. Other habitat variables were poor predictors of species richness. The lack of a relationship between habitat and species richness appeared to be a consequence of most vegetation types present not being sufficiently distinct for birds to differentially associate with them. I also found no relationship between bird population density and area, suggesting that habitat quality in marshes did not improve with increasing size, and species evenness declined with increasing richness because greater richness was associated with the presence of more rare species. Larger marshes had more rare species, species with larger populations, and species with a minimum threshold area for occurrence. Thus, my results are consistent with theoretical predictions that larger populations are less prone to local extinction and, as individuals are added to a community, more rare species are present.     

Avian fitness consequences match habitat selection at the nest‐site and landscape scale in agriculturally fragmented landscapes

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人工巢箱繁殖鸟类主要巢捕食者及其影响因素

巢捕食是影响鸟类繁殖成功率的一个重要因素,也是鸟类繁殖生态研究中的一项重要内容。确定鸟类的主要巢捕食者及影响巢捕食的因素对于了解鸟类繁殖成功率、种群增长率及种群数量等具有重要意义。2009—2012年,对辽宁仙人洞国家级自然保护区人工巢箱中繁殖的杂色山雀(Parus varius)、沼泽山雀(P.palustris)、大山雀(P.major)和白眉姬鹟(Ficedula zanthopygia)四种鸟类的巢捕食率及影响巢捕食的因素进行了研究。研究共记录到238个繁殖巢(杂色山雀74巢、沼泽山雀21巢、大山雀118巢、白眉姬鹟25巢),其中35巢被捕食,捕食率为14.7%,雏鸟期被捕食占91.4%。巢捕食率在4种鸟类之间无差异(x~2=0.429,df=3,P=0.934)。以锦蛇(Elaphe spp.)为代表的蛇类是该地区小型森林洞巣鸟类繁殖期主要捕食者,占总捕食率的94.3%。对影响巢捕食的22个相关因子进行二元逻辑斯蒂回归分析发现,坡度、地面裸露率、草本盖度对巢捕食具有显著性影响;出雏时间、坡位、距碎石块距离对巢捕食的影响接近显著水平;而巢高、树粗、巢箱年龄、窝卵数、距路距离等对巢捕食无显著影响。因此,处于坡度较陡,坡位较高,草本覆盖率较高,地面裸露率较低,距碎石块距离较近,且出雏时间较晚的巢更容易被捕食。     

Rapid climate‐driven loss of breeding habitat for Arctic migratory birds

Millions of birds migrate to and from the Arctic each year, but rapid climate change in the High North could strongly affect where species are able to breed, disrupting migratory connections globally. We modelled the climatically suitable breeding conditions of 24 Arctic specialist shorebirds and projected them to 2070 and to the mid‐Holocene climatic optimum, the world's last major warming event ~6000 years ago. We show that climatically suitable breeding conditions could shift, contract and decline over the next 70 years, with 66–83% of species losing the majority of currently suitable area. This exceeds, in rate and magnitude, the impact of the mid‐Holocene climatic optimum. Suitable climatic conditions are predicted to decline acutely in the most species rich region, Beringia (western Alaska and eastern Russia), and become concentrated in the Eurasian and Canadian Arctic islands. These predicted spatial shifts of breeding grounds could affect the species composition of the world's major flyways. Encouragingly, protected area coverage of current and future climatically suitable breeding conditions generally meets target levels; however, there is a lack of protected areas within the Canadian Arctic where resource exploitation is a growing threat. Given that already there are rapid declines of many populations of Arctic migratory birds, our results emphasize the urgency of mitigating climate change and protecting Arctic biodiversity.     

Fitness consequences of timing of breeding in birds: date effects in the course of a reproductive episode

In seasonal environments, avian reproductive performance almost generally declines in the course of the season. Quantifying the associated fitness consequences of timing of breeding, i.e. of date‐related factors, is important for understanding the evolution of temporal patterns in avian life‐histories and for predicting consequences of climate change. The seasonal decline can also be caused by an effect of parental quality: individuals with high phenotypic quality may breed early. The results of existing experimental studies investigating whether date or quality effects cause the seasonal decline are inconsistent, indicating that both mechanisms might be involved. However, it remains unclear to what extent the confounding effect of quality occurs and what the fitness consequences of timing per se over a whole breeding episode are. In a cross‐fostering experiment using the barn swallows’ second broods we evaluated the causes for the seasonal decline in reproductive performance for three distinct periods of a reproductive attempt, the early nestling period, the late nestling period and the post‐fledging period, and we assessed the overall fitness consequences of timing per se. A seasonal decline in juvenile feather growth rate was mainly due to date effects in the late nestling period, although we determined quality effects during early nestling development. Date effects on survival were present in the post‐fledging period, but not in the nestling period. The decline in feather length due to date effects in the nestling period accounted for 9% of the seasonal decline in post‐fledging survival, whereas date effects arising only in the post‐fledging period caused 91% of the decline. These results suggest that date effects increase in the course of a reproductive episode. Thus, the benefits of an early timing of breeding can be quantified only when considering also the post‐fledging period. We suggest that the timing of breeding evolved through a trade‐off between date‐related benefits and quality‐related costs of early breeding.     

Nutrient limitation in species‐rich Calthion grasslands in relation to opportunities for restoration in a peat meadow landscape

Questions: Which nutrient(s) limit(s) vegetation productivity in Calthion grasslands? Is phosphorus release a bottleneck for restoration of species‐rich Calthion grasslands on rewetted dairy meadows? Location: Three species‐rich Calthion grasslands in the Western Peat District in the Netherlands. Methods: We conducted a field fertilization experiment with nitrogen (N), phosphorus (P) and potassium (K) in three existing Calthion grasslands to evaluate the potential for restoration on rewetted dairy meadows. Responses of above‐ground biomass, tissue nutrient concentrations and nutrient ratios were determined after 2 yr of fertilization. Results: Biomass increased with fertilization with N‐only and K‐only but did not react to P‐only additions. Comparisons of tissue nutrient concentrations and nutrient ratios also gave indications of N and K limitation. Conclusions: The strong P release expected after rewetting should not necessarily interfere with restoration of Calthion communities on rewetted dairy meadows. It is concluded that for successful restoration management measures should focus on reducing N and/or K availability. Potassium might be an overlooked bottleneck in the restoration of species‐rich grasslands.     

Species‐area relationships of lemurs in a fragmented landscape in Madagascar

We used species‐area relationships (SARs) to investigate the effects of habitat loss on lemur biogeography. We measured species richness via visual surveys on line transects within 42 fragments of dry deciduous forest at the Ambanjabe field site in Ankarafantsika National Park, Madagascar. We measured human disturbance and habitat characteristics within 38 of the 42 fragments. We measured the distance between each fragment and the nearest settlement, continuous forest, and nearest neighboring fragment. We fit 10 candidate SAR models to the data using nonlinear least squares regression and compared them using Akaike's Information Criterion (AIC). To determine how habitat characteristics, as well as area, influenced species richness, we ran a hierarchical partitioning procedure to select which variables to include in generalized additive models (GAMs) and compared them using AIC. Contrary to expectations, we found that lemurs form convex SARs, without a “small island effect”, and with the power model being the most likely SAR model. Although we found that four variables (area, survey effort, and total human disturbance, and mean tree height) independently contributed greater than 10% of the variation in lemur species richness, only area was included in the most likely model. We suggest that the power model was the most likely SAR model and our inability to detect a “small island effect” are the result of Microcebus spp. being edge tolerant and capable of dispersing through matrix, scale issues in the study design, and low γ‐diversity in the landscape. However, more study is needed to determine what role human disturbance plays in influencing species richness in lemurs.     

Behavioral signature of intraspecific competition and density dependence in colony‐breeding marine predators

In populations of colony‐breeding marine animals, foraging around colonies can lead to intraspecific competition. This competition affects individual foraging behavior and can cause density‐dependent population growth. Where behavioral data are available, it may be possible to infer the mechanism of intraspecific competition. If these mechanics are understood, they can be used to predict the population‐level functional response resulting from the competition. Using satellite relocation and dive data, we studied the use of space and foraging behavior of juvenile and adult gray seals (Halichoerus grypus) from a large (over 200,000) and growing population breeding at Sable Island, Nova Scotia (44.0 ^oN 60.0 ^oW). These data were first analyzed using a behaviorally switching state‐space model to infer foraging areas followed by randomization analysis of foraging region overlap of competing age classes. Patterns of habitat use and behavioral time budgets indicate that young‐of‐year juveniles (YOY) were likely displaced from foraging areas near (<10 km) the breeding colony by adult females. This displacement was most pronounced in the summer. Additionally, our data suggest that YOY are less capable divers than adults and this limits the habitat available to them. However, other segregating mechanisms cannot be ruled out, and we discuss several alternate hypotheses. Mark–resight data indicate juveniles born between 1998 and 2002 have much reduced survivorship compared with cohorts born in the late 1980s, while adult survivorship has remained steady. Combined with behavioral observations, our data suggest YOY are losing an intraspecific competition between adults and juveniles, resulting in the currently observed decelerating logistic population growth. Competition theory predicts that intraspecific competition resulting in a clear losing competitor should cause compensatory population regulation. This functional response produces a smooth logistic growth curve as carrying capacity is approached, and is consistent with census data collected from this population over the past 50 years. The competitive mechanism causing compensatory regulation likely stems from the capital‐breeding life‐history strategy employed by gray seals. This strategy decouples reproductive success from resources available around breeding colonies and prevents females from competing with each other while young are dependent.     

Sex‐biased breeding dispersal is predicted by social environment in birds

Sex‐biased dispersal is common in vertebrates, although the ecological and evolutionary causes of sex differences in dispersal are debated. Here, we investigate sex differences in both natal and breeding dispersal distances using a large dataset on birds including 86 species from 41 families. Using phylogenetic comparative analyses, we investigate whether sex‐biased natal and breeding dispersal are associated with sexual selection, parental sex roles, adult sex ratio (ASR), or adult mortality. We show that neither the intensity of sexual selection, nor the extent of sex bias in parental care was associated with sex‐biased natal or breeding dispersal. However, breeding dispersal was related to the social environment since male‐biased ASRs were associated with female‐biased breeding dispersal. Male‐biased ASRs were associated with female‐biased breeding dispersal. Sex bias in adult mortality was not consistently related to sex‐biased breeding dispersal. These results may indicate that the rare sex has a stronger tendency to disperse in order to find new mating opportunities. Alternatively, higher mortality of the more dispersive sex could account for biased ASRs, although our results do not give a strong support to this explanation. Whichever is the case, our findings improve our understanding of the causes and consequences of sex‐biased dispersal. Since the direction of causality is not yet known, we call for future studies to identify the causal relationships linking mortality, dispersal, and ASR.     

Exploring co‐occurrence of closely‐related guild members in a fragmented landscape subject to rapid transformation

Quantifying biotic responses to landscape transformation is a major research focus. Most past studies have explored co‐occurrence of entire communities of a given group (e.g. birds) within largely intact ecosystems or over a limited time‐frame. By contrast, here we use data from a 15 yr experimental study, to explore intra‐guild co‐occurrence of six closely‐related and functionally‐similar sets of birds within 55 woodland fragments. Areas surrounding these remnants are undergoing transformation from grazed paddocks to Pinus radiata plantations, leading to a novel assemblage of forest and woodland birds. We sought to determine if the occurrence of a given species in a guild influenced the occurrence of other closely‐related species in that guild, and through this relationship whether there was evidence of co‐occurrence between species. After controlling for environmental and habitat variables which can affect species occurrence like time since commencement of landscape transformation, patch size and vegetation type, we found the occurrence of a given species was influenced by the occurrence of a closely‐related species in the same guild. Co‐occurrence varied among bird guilds and included: 1) positive co‐occurrence in which occurrence of one species within fragments positively affected the occurrence of another closely‐related guild member (e.g. eastern and crimson rosellas); and 2) negative co‐occurrence in which the occurrence of one species was negatively associated with the occurrence of another within the same guild (e.g. willie wagtail and grey fantail). We also identified interactions between patch size and species recording frequency within members of two guilds. For example, modelling of conditional recording frequency revealed the eastern rosella increased with increasing recordings of the crimson rosella in large patches, but decreased with increasing recordings of the crimson rosella in small patches. Our results provide empirical evidence of co‐occurrence among guild members and underscore the complexity of biotic responses to landscape transformation.     

Minimum area requirements of breeding birds in fragmented woodland of Central Argentina

The breeding avifauna of 25 woodland fragments (0.85–280 ha) was studied between 1996 and 2004 in Córdoba, Argentina. A distinctive feature of the avifauna of the fragments studied is the low area requirement of most of the species. Of the 54 woodland species recorded, 32 (59.3%) require c . 1 ha and 43 (79.6%) needed no more than 3 ha. Also noticeable is the relatively high number of individuals of most of the species. Both characteristics suggest a good tolerance to fragmentation. However, nine species (16.7%), the area-sensitive species, need fragments of 80 ha or larger. Moreover, eight resident species have apparently become extinct in the fragments that were studied, including the five large species that originally inhabited the area. Proportional odds and log linear models were fitted to relate the minimum area requirements of these species to various ecological characteristics (body size, diet, habitat use, migratory status, and nest type). Closed nesters, resident, woodland exterior, and medium-size species were the groups requiring larger areas. No apparent pattern was observed between area requirements and diet. A recent law prohibits woodland removal, which is a positive step towards the conservation of the fragments. The results suggest that as long as the present amount of woodland is maintained, the situation of the avifauna of the fragments is relatively secure, at least in the short term.     

Changes in Danish farmland habitats and their populations of breeding birds

Habitat changes and the breeding bird populations were studied at Kraghede, N Jutland the years 1970–1980. Generally, agricultural practice was intensified leading to an increase in the area used permanently for growing barley, and the abandonment of grasslands and meadows. Animal husbandry decreased, until it was restricted to a small number of farms.
These habitat changes were the largest ones to occur in the Danish landscape for some 150–200 yr and had a significant influence on the breeding birds. Species confined to the meadows, such as the lapwing Vanellus vanellus , meadow pipit Anthus pratensis and others, decreased by 50–80% or even more. Species associated with the grassland when feeding, like the swallow Hirundo rustica , starling Sturnns vulgaris and magpie Pica pica , all had lower population levels at the end of the 1970's compared to those at the beginning of the decade.     

Species traits and the response of open‐habitat species to forest edge in landscape mosaics

Human driven changes in land‐use have increased the need to understand how landscape structure affects species distribution. We studied how forest edges affected the distribution of birds in grasslands recently encroached by forest patches. We investigated how species’ biological traits influenced their response to vegetation change near forest edges. We censured birds along 300‐m line transects run into the open habitat perpendicularly to forest edges. We recorded habitat variables and landscape context along each transect and characterized edges and forest patches. We recorded 33 bird species in 153 transects for a total of 654 individuals. We analyzed species response to edges with generalized linear mixed models. Habitat preference was prevalent to explain species response to forest edges. The abundance of open‐habitat birds such as skylark Alauda arvensis decreased significantly in the vicinity of edges. This negative response extended within 150 m from the edge. The effect was disproportionately higher in open‐habitat species with high conservation concern. The abundance of species feeding or/and breeding in both forest and open habitat, such as woodlarks Lullula arborea, sharply increased near edges (positive edge response). Abundance of shrub and non‐shrub dependent species increased with distance to edge. The two species groups did no differ in abundance/distance to edge relationship. Intensity of species response to forest edges varied among transects in relation to transect vegetation characteristics. Edge length or aspect, diet and nest height had no direct effect. We discuss the possible role of variation in resources and nest predation risk to explain observed patterns.     

Levels of antioxidants in rural and urban birds and their consequences

Numerous animals have successfully invaded urban habitats, although the factors associated with invasion success remain poorly understood. Urban areas are characterized by warmer microclimates, higher levels of primary productivity, longer breeding seasons and higher levels of pollutants. All these factors should cause oxidative stress, favoring invasion by species that have access to high levels of antioxidants. We analyzed concentrations of two categories of dietary, fat-soluble antioxidants (total carotenoids, total vitamin E) in the liver, the main storage organ in birds. Individuals killed by cats had lower levels of vitamin E than individuals that died for other reasons, showing natural selection on stored antioxidants. Bird species that had successfully colonized urban areas had significantly higher levels of vitamin E and total carotenoids than species that did not succeed, and rural populations had higher concentrations of vitamin E and total carotenoids than urban populations of the same species. Interspecific differences in concentrations of fat-soluble antioxidants, and differences between rural and urban populations of the same species, were accounted for by diet, but also by time since urbanization and number of generations since urbanization. These findings suggest that antioxidants, and by implication the ability to cope with oxidative stress, have contributed to successful invasion of urban areas by birds, and that the concentration of these antioxidants has changed in response to the urban environment.