Ecological filtering shapes the impacts of agricultural deforestation on biodiversity.

The biodiversity impacts of agricultural deforestation vary widely across regions. Previous efforts to explain this variation have focused exclusively on the landscape features and management regimes of agricultural systems, neglecting the potentially critical role of ecological filtering in shaping deforestation tolerance of extant species assemblages at large geographical scales via selection for functional traits. Here we provide a large-scale test of this role using a global database of species abundance ratios between matched agricultural and native forest sites that comprises 71 avian assemblages reported in 44 primary studies, and a companion database of 10 functional traits for all 2,647 species involved. Using meta-analytic, phylogenetic and multivariate methods, we show that beyond agricultural features, filtering by the extent of natural environmental variability and the severity of historical anthropogenic deforestation shapes the varying deforestation impacts across species assemblages. For assemblages under greater environmental variability-proxied by drier and more seasonal climates under a greater disturbance regime-and longer deforestation histories, filtering has attenuated the negative impacts of current deforestation by selecting for functional traits linked to stronger deforestation tolerance. Our study provides a previously largely missing piece of knowledge in understanding and managing the biodiversity consequences of deforestation by agricultural deforestation.

Bird tolerance to humans in open tropical ecosystems.

Animal tolerance towards humans can be a key factor facilitating wildlife-human coexistence, yet traits predicting its direction and magnitude across tropical animals are poorly known. Using 10,249 observations for 842 bird species inhabiting open tropical ecosystems in Africa, South America, and Australia, we find that avian tolerance towards humans was lower (i.e., escape distance was longer) in rural rather than urban populations and in populations exposed to lower human disturbance (measured as human footprint index). In addition, larger species and species with larger clutches and enhanced flight ability are less tolerant to human approaches and escape distances increase when birds were approached during the wet season compared to the dry season and from longer starting distances. Identification of key factors affecting animal tolerance towards humans across large spatial and taxonomic scales may help us to better understand and predict the patterns of species distributions in the Anthropocene.

Common Whitethroats Curruca communis show a continuum of residency duration but a high degree of between-years site fidelity at nonbreeding grounds in Nigeria.

The nonbreeding period represents a significant part of an Afro-Palearctic migratory bird's annual cycle. Decisions such as whether to remain at a single site and whether to return to it across years have important effects on aspects such as survival, future breeding success, migratory connectivity, and conservation. During this study, we color-ringed 337 common Whitethroats Curruca communis and undertook daily resightings to understand site persistence and the degree of site fidelity throughout three nonbreeding periods (November-April) in Nigeria. The probability of detecting a color-ringed Whitethroat when it was present was 0.33. Site persistence varied widely across individuals (1-165 days) and did not differ significantly with sex or year, though first-year birds remained for significantly shorter periods than adults. We believe that shorter residencies are likely due to the use of multiple stationary nonbreeding sites rather than low winter survival. A minimum of 19% of individuals returned to the study site the following year and shifted, on average, 300 m, suggesting that Whitethroats have a relatively high degree of between-years site fidelity at a very fine scale. An individual's previous residency duration did not seem to determine its residency duration the following year. We suggest that spatial fidelity is high and constant through years, but temporal fidelity is not, and individual residency patterns vary, probably according to yearly and seasonal conditions. Our results highlight the complexity of the annual cycle of a single species and the importance of carrying out in situ, fine-scale research throughout a migrant's annual cycle over several years.

Age-related changes in migratory behaviour within the first annual cycle of a passerine bird.

First time migrants (juveniles hereafter) of many species migrate without specific knowledge of non-breeding locations, but experience may aid adults in timing and route decisions because they can migrate more efficiently to their previous non-breeding sites. Consequently, we expect a transition to more efficient migratory behaviour with age, but when and how this happens is little known. We used light-level geolocator data from Cyprus wheatears Oenanthe cypriaca to compare migration timing and route directness between juveniles and adults, and repeatability of their timing and non-breeding locations. We predicted that juveniles would depart and arrive later than adults for both autumn and spring migration; that duration of migration would be greater for juveniles; that routes taken by juveniles would be less direct than those for adults; and that autumn and spring departure timing, and non-breeding locations, would be more repeatable for adults between two years than for juveniles between their first and subsequent migration. We found that juveniles departed significantly later than adults in autumn but there was no difference in arrival timing, and although spring departure timings did not differ, juveniles arrived on the breeding grounds later than adults. Nevertheless, we found no significant age-related difference in the duration of migration in autumn or spring. Yet, juvenile migrations were less direct than those of adults in autumn, but not spring. We found evidence that spring departure timing and non-breeding locations were repeatable for adults but not juveniles. Our findings show that age-related changes in migratory behaviour begin to occur during the first annual cycle demonstrating the potential for early adaptation to environmental variability within an individual's life.

Migratory routes, breeding locations and multiple non-breeding sites of Common Whitethroats Curruca communis revealed by geolocators.

Understanding general migration characteristics and how breeding and non-breeding sites are connected is crucial for predicting the response of long-distance migratory bird populations to environmental changes. We use data collected from six geolocators to describe migratory routes and identify breeding and non-breeding locations, migratory behaviour and differences between spring and autumn migration of Common Whitethroats Curruca communis, an Afro-Palearctic migrant, wintering in Nigeria. Most individuals departed on spring migration in April, following a north-easterly direction, arriving at their breeding grounds across central-eastern Europe (~425,000 km2) in May. Departures from breeding grounds took place between July and August in a south-westerly direction. During spring migration individuals travelled longer distances at faster rates making its overall duration shorter than autumn migration. We suggest that, while Whitethroats can cross the Sahara Desert and Mediterranean Sea in a single flight, they are likely to refuel before and after crossing. Results indicate that Whitethroats undertook loop migration and visited two wintering sites: first in the Sahel, then in Nigeria, where they remained until spring migration. Geolocator results and data from the European Union for Bird Migration's (EURING) ringing database suggest that Whitethroats have a relatively high migratory spread-individuals from a single non-breeding site breed across a wide area of Europe. Our research is the first to track and describe the complete annual cycle of Whitethroats and one of the few studies to do so for any Afro-Palearctic migrant from non-breeding grounds. We identified the Sahel as an important refuelling and first wintering site indicating its conservation, alongside other stopover sites, is crucial for the species. We believe that changes in this region will have severe effects on a subset of individuals of specific European breeding populations, but these effects will greatly depend on the severity of the changes and at what spatial scale they occur.

The potential function of post-fledging dispersal behavior in first breeding territory selection for males of a migratory bird.

One possible hypothesis for the function of post-fledging dispersal is to locate a suitable future breeding area. This post-fledging period may be particularly important in migratory species because they have a limited period to gather information prior to autumn migration, and in protandrous species, males must quickly acquire a territory after returning from spring migration to maximize their fitness. Here we use color-ring resightings to investigate how the post-fledging dispersal movements of the Cyprus wheatear Oenanthe cypriaca, a small migratory passerine, relate to their first breeding territory the following year when they return from migration. We found that males established first breeding territories that were significantly closer to their post-fledging location than to their natal sites or to post-fledging locations of other conspecifics, but these patterns were not apparent in females. Our findings suggest that familiarity with potential breeding sites may be important for juveniles of migratory species, particularly for the sex that acquires and advertises breeding territories. Exploratory dispersal prior to a migrant's first autumn migration may contribute toward its breeding success the following year, further highlighting the importance of early seasonal breeding on fitness and population dynamics more generally.

Pest-removal services provided by birds on subsistence farms in south-eastern Nigeria.

To what extent birds provide the ecosystem service of pest control in subsistence farms, and how this service might depend on retained natural habitats near farmlands is unexplored in West Africa. To fill this knowledge gap, we placed plasticine mimics of insect pests on experimentally grown crops on the Mambilla Plateau, South Eastern Nigeria. We recorded bird attacks on the mimics and the proportion of mimics removed by birds. We also determined the influence of distance of crops from forest fragments on both attack and removal rates. We placed 90 potted plants of groundnut (Arachis hypogea) and bambara nut (Vigna subterranea) along 15 transects running 4.5 km from forest edge into open grassland. Each plant had six of the 540 mimics in total placed on their leaves. We inspected the potted plants weekly for 12 weeks to record (i) the presence of bird beak marks on mimics, and (ii) the number of missing mimics. Once a week we collected all the mimics from the plants and counted the number of assumed beak marks. After counting we replaced the mimics on the plants, mark free. We found a strong positive correlation between the abundance of insectivorous birds and the mean number of missing mimics and/or bird attack marks on mimics. However, this positive effect of insectivorous bird abundance on prey mimic attack/removal became less strong the farther they were from a forest fragment. We found increased predation rates and abundance of insectivorous birds closer to forest fragments. Our data suggest that pest predation may be a key ecosystem service provided by insectivorous birds on Nigerian farmlands. Farmlands that are closer to forest fragments may experience a higher rate of pest control by insectivorous birds than those further away, suggesting that retaining forest fragments in the landscape may enhance pest control services in sub-Saharan subsistence farms.

Geographic variation in baseline innate immune function does not follow variation in aridity along a tropical environmental gradient.

Geographic variation in aridity determines environmental productivity patterns, including large-scale variability in pathogens, vectors and associated diseases. If disease risk decreases with increasing aridity and is matched by immune defense, we predict a decrease in innate immune function along a gradient of increasing aridity from the cool-wet forest to the hot-dry Sahel, from south to north in Nigeria. We sampled blood and measured five innate immune indices from 286 Common Bulbuls Pycnonotus barbatus between 6 and 13°N. We sampled in the dry season; we resampled the first location (Jos) also as the last sample location to test temporal change in immune function. Immune indices did not decrease with aridity. One immune index, nitric oxide concentration showed a weak quadratic pattern. In Jos, ovotransferrin concentration, haemagglutination and haemolysis titres increased 12 weeks into the dry season, contrary to expectations that immune indices should decrease with increased dryness. In this tropical system, innate immune function does not decrease with increasing aridity but temporal factors within a location may influence immune function more strongly than spatial variation in aridity, suggesting that immune variation does not follow a simple environmental productivity pattern. Consequently, caution should probably be exercised in predicting effects of climate variability on immune function or disease risk.

A fruit diet rather than invertebrate diet maintains a robust innate immunity in an omnivorous tropical songbird.

Diet alteration may lead to nutrient limitations even in the absence of food limitation, and this may affect physiological functions, including immunity. Nutrient limitations may also affect the maintenance of body mass and key life-history events that may affect immune function. Yet, variation in immune function is largely attributed to energetic trade-offs rather than specific nutrient constraints. To test the effect of diet on life-history traits, we tested how diet composition affects innate immune function, body mass and moult separately and in combination with each other, and then used path analyses to generate hypotheses about the mechanistic connections between immunity and body mass under different diet compositions. We performed a balanced parallel and crossover design experiment with omnivorous common bulbuls Pycnonotus barbatus in out-door aviaries in Nigeria. We fed 40 wild-caught bulbuls ad libitum on fruits or invertebrates for 24 weeks, switching half of each group between treatments after 12 weeks. We assessed innate immune indices (haptoglobin, nitric oxide and ovotransferrin concentrations, and haemagglutination and haemolysis titres), body mass and primary moult, fortnightly. We simplified immune indices into three principal components (PCs), but we explored mechanistic connections between diet, body mass and each immune index separately. Fruit-fed bulbuls had higher body mass, earlier moult and showed higher values for two of the three immune PCs compared to invertebrate-fed bulbuls. These effects were reversed when we switched bulbuls between treatments after 12 weeks. Exploring the correlations between immune function, body mass and moult, showed that an increase in immune function was associated with a decrease in body mass and delayed moult in invertebrate-fed bulbuls, while fruit-fed bulbuls maintained body mass despite variation in immune function. Path analyses indicated that diet composition was most likely to affect body mass and immune indices directly and independently from each other. Only haptoglobin concentration was indirectly linked to diet composition via body mass. We demonstrated a causal effect of diet composition on innate immune function, body mass and moult: bulbuls were in a better condition when fed on fruits than invertebrates, confirming that innate immunity is nutrient specific. Our results are unique because they show a reversible effect of diet composition on wild adult birds whose immune systems are presumably fully developed and adapted to wild conditions-demonstrating a short-term consequence of diet alteration on life-history traits.

FragSAD: A database of diversity and species abundance distributions from habitat fragments.

Habitat destruction is the single greatest anthropogenic threat to biodiversity. Decades of research on this issue have led to the accumulation of hundreds of data sets comparing species assemblages in larger, intact, habitats to smaller, more fragmented, habitats. Despite this, little synthesis or consensus has been achieved, primarily because of non-standardized sampling methodology and analyses of notoriously scale-dependent response variables (i.e., species richness). To be able to compare and contrast the results of habitat fragmentation on species' assemblages, it is necessary to have the underlying data on species abundances and sampling intensity, so that standardization can be achieved. To accomplish this, we systematically searched the literature for studies where abundances of species in assemblages (of any taxa) were sampled from many habitat patches that varied in size. From these, we extracted data from several studies, and contacted authors of studies where appropriate data were collected but not published, giving us 117 studies that compared species assemblages among habitat fragments that varied in area. Less than one-half (41) of studies came from tropical forests of Central and South America, but there were many studies from temperate forests and grasslands from all continents except Antarctica. Fifty-four of the studies were on invertebrates (mostly insects), but there were several studies on plants (15), birds (16), mammals (19), and reptiles and amphibians (13). We also collected qualitative information on the length of time since fragmentation. With data on total and relative abundances (and identities) of species, sampling effort, and affiliated meta-data about the study sites, these data can be used to more definitively test hypotheses about the role of habitat fragmentation in altering patterns of biodiversity. There are no copyright restrictions. Please cite this data paper and the associated Dryad data set if the data are used in publications.

Seasonal differences in baseline innate immune function are better explained by environment than annual cycle stage in a year-round breeding tropical songbird.

Seasonal variation in innate immunity is often attributed to either temporal environmental variation or to life-history trade-offs that arise from specific annual cycle stages but decoupling them is difficult in natural populations. Here, we effectively decouple seasonal environmental variation from annual cycle stage effects by exploiting cross-seasonal breeding and moult in the tropical Common Bulbul Pycnonotus barbatus. We test how annual cycle stage interacts with a key seasonal environmental variable, rainfall, to determine immunity at population and individual level. If immune challenge varies with precipitation, we might expect immune function to be higher in the wet season due to increased environmental productivity. If breeding or moult imposes resource constraints on birds, depending on or independent of precipitation, we might expect lower immune indices during breeding or moult. We sampled blood from 818 birds in four annual cycle stage categories: breeding, moult, simultaneous breeding and moulting, or neither. We quantified indices of innate immunity (haptoglobin, nitric oxide (NOx ) and ovotransferrin concentrations, and haemagglutination and haemolysis titres) over two annual cycles of wet and dry seasons. Environment (but not annual cycle stage or interactions between both) explained variation in all immune indices, except NOx . NOx concentration differed between annual cycle stages but not between seasons. However, within the wet season, haptoglobin, NOx , ovotransferrin and haemolysis differed significantly between breeding and non-breeding females. Aside from some recorded inconsistencies, population level results were largely similar to results within individuals that were measured repeatedly. Unexpectedly, most immune indices were higher in the dry season and during breeding. Higher immune indices may be explained if fewer or poorer quality resources force birds to increase social contact, thereby exposing individuals to novel antigens and increased infection risk, independently of environmental productivity. Breeding birds may also show higher immunity if less immune-competent and/or infected females omit breeding. We conclude that seasonal environmental variation impacts immunity more directly in natural animal populations than via resource trade-offs. In addition, immune indices were more often variable within than among individuals, but some indices are characteristic of individuals, and so may offer selective advantages if heritable.

Population consequences of migratory variability differ between flyways.

Long-distance migratory bird populations are likely to be declining because of climate change shifting habitats or anthropogenic habitat loss [1], but this may be mediated by the size of the non-breeding area over which a population spreads (migratory spread), and migration distance (or number of stop-over sites). High migratory spread may make populations more resilient to climate change because they already encompass shifting habitats, but less resilient to uneven habitat loss that may not affect populations with low migratory spread [2] (Figure 1C). As migration distance increases so the probability of encountering a stop-over site with negative environmental change increases [3] (Figure 1D). Consequently, if habitat shift through climate change is the main driver of declines we predict more positive population trends for high spread migrants, but the reverse for outright habitat loss (Figure 1E); we also predict negative population trends for longer distance migrants (Figure 1F). But these relationships may vary between flyways, which differ profoundly in their climate variation, human population change and geography. Here we show that climate change may be more important in Neotropic migrant population declines whereas habitat loss may be more important in the Afro-Palearctic.

Temperature and aridity determine body size conformity to Bergmann's rule independent of latitudinal differences in a tropical environment.

Bergmann's rule, defined as the tendency for endotherms to be larger in colder environments, is a biophysical generalization of body size variation that is frequently tested along latitudinal gradients, even though latitude is only a proxy for temperature variation. We test whether variation in temperature and aridity determine avian body size conformity to Bergmann's rule independent of latitude differences, using the ubiquitous Common Bulbul Pycnonotus barbatus, along a West African environmental gradient. We trapped 538 birds in 22 locations between latitudes 6 and 13°N in Nigeria, and estimated average body surface area to mass ratio per location. We then modelled body surface to mass ratio using general linear models, with latitude, altitude and one of 19 bioclimatic variables extracted from http://www.worldclim.org/bioclim as predictors. We sequentially dropped latitude and altitude from each model to obtain the R 2 of the resultant models. Finally, we compared the R 2 of univariate models, where bioclimatic variables predicted body surface area to mass ratio significantly (14 out of 19), to multivariate models including latitude, altitude and a bioclimatic variable, using the Wilcoxon matched pairs test. We found that multivariate models did not perform better than univariate models with only bioclimatic variables. Six temperature and eight precipitation variables significantly predicted variation in body surface area to mass ratio between locations; in fact, 50% (seven out of 14) of these better explained variation in body surface area to mass ratio than the multivariate models. Birds showed a larger body surface area relative to body mass ratio in hotter environments independent of latitude or altitude, which conforms to Bergmann's rule. Yet, a combination of morphometric analyses and controlled temperature-exposure experiments is required to prove the proposed relationship between relative body surface area and thermoregulation in endotherms.

User fees across ecosystem boundaries: Are SCUBA divers willing to pay for terrestrial biodiversity conservation?

While ecological links between ecosystems have been long recognised, management rarely crosses ecosystem boundaries. Coral reefs are susceptible to damage through terrestrial run-off, and failing to account for this within management threatens reef protection. In order to quantify the extent to that coral reef users are willing to support management actions to improve ecosystem quality, we conducted a choice experiment with SCUBA divers on the island of Bonaire, Caribbean Netherlands. Specifically, we estimated their willingness to pay to reduce terrestrial overgrazing as a means to improve reef health. Willingness to pay was estimated using the multinomial, random parameter and latent class logit models. Willingness to pay for improvements to reef quality was positive for the majority of respondents. Estimates from the latent class model determined willingness to pay for reef improvements of between $31.17 - $413.18/year, dependent on class membership. This represents a significant source of funding for terrestrial conservation, and illustrates the potential for user fees to be applied across ecosystem boundaries. We argue that such across-ecosystem-boundary funding mechanisms are an important avenue for future investigation in many connected systems.

Low migratory connectivity is common in long-distance migrant birds.

Estimating how much long-distance migrant populations spread out and mix during the non-breeding season (migratory connectivity) is essential for understanding and predicting population dynamics in the face of global change. We quantify variation in population spread and inter-population mixing in long-distance, terrestrial migrant land-bird populations (712 individuals from 98 populations of 45 species, from tagging studies in the Neotropic and Afro-Palearctic flyways). We evaluate the Mantel test as a metric of migratory connectivity, and explore the extent to which variance in population spread can be explained simply by geography. The mean distance between two individuals from the same population during the non-breeding season was 743 km, covering 10-20% of the maximum width of Africa/South America. Individuals from different breeding populations tended to mix during the non-breeding season, although spatial segregation was maintained in species with relatively large non-breeding ranges (and, to a lesser extent, those with low population-level spread). A substantial amount of between-population variation in population spread was predicted simply by geography, with populations using non-breeding zones with limited land availability (e.g. Central America compared to South America) showing lower population spread. The high levels of population spread suggest that deterministic migration tactics are not generally adaptive; this makes sense in the context of the recent evolution of the systems, and the spatial and temporal unpredictability of non-breeding habitat. The conservation implications of generally low connectivity are that the loss (or protection) of any non-breeding site will have a diffuse but widespread effect on many breeding populations. Although low connectivity should engender population resilience to shifts in habitat (e.g. due to climate change), we suggest it may increase susceptibility to habitat loss. We hypothesize that, because a migrant species cannot adapt to both simultaneously, migrants generally may be more susceptible to population declines in the face of concurrent anthropogenic habitat and climate change.

Mushroom tyrosinase: A model system to combine experimental investigation of enzyme-catalyzed reactions, data handling using R, and enzyme-inhibitor structural studies.

The activity of mushroom tyrosinase can be measured by monitoring the conversion of phenolic compounds into quinone derivatives using spectrophotometry. This article describes a series of experiments which characterize the functional properties of tyrosinase, the analysis of the resulting data using R to determine the kinetic parameters, and the exploration of the structural properties of tyrosinase-inhibitor complexes. Tyrosinase assay development and subsequent activity measurements, in the presence of varying pH, substrate concentration and inhibitors, offers the opportunity to learn the enzyme characterization skills relevant to a research laboratory setting. Combining the activity studies with an exploration of the nature of the tyrosinase-inhibitor interactions enables a structural understanding of the experimental observations.

Biparental incubation-scheduling: no experimental evidence for major energetic constraints.

Incubation is energetically demanding, but it is debated whether these demands constrain incubation-scheduling (i.e., the length, constancy, and timing of incubation bouts) in cases where both parents incubate. Using 2 methods, we experimentally reduced the energetic demands of incubation in the semipalmated sandpiper, a biparental shorebird breeding in the harsh conditions of the high Arctic. First, we decreased the demands of incubation for 1 parent only by exchanging 1 of the 4 eggs for an artificial egg that heated up when the focal bird incubated. Second, we reanalyzed the data from the only published experimental study that has explicitly tested energetic constraints on incubation-scheduling in a biparentally incubating species (Cresswell et al. 2003). In this experiment, the energetic demands of incubation were decreased for both parents by insulating the nest cup. We expected that the treated birds, in both experiments, would change the length of their incubation bouts, if biparental incubation-scheduling is energetically constrained. However, we found no evidence that heating or insulation of the nest affected the length of incubation bouts: the combined effect of both experiments was an increase in bout length of 3.6min (95% CI: -33 to 40), which is equivalent to a 0.5% increase in the length of the average incubation bout. These results demonstrate that the observed biparental incubation-scheduling in semipalmated sandpipers is not primarily driven by energetic constraints and therefore by the state of the incubating bird, implying that we still do not understand the factors driving biparental incubation-scheduling.

Predicting population-level risk effects of predation from the responses of individuals.

Fear of predation produces large effects on prey population dynamics through indirect risk effects that can cause even greater impacts than direct predation mortality. As yet, there is no general theoretical framework for predicting when and how these population risk effects will arise in specific prey populations, meaning that there is often little consideration given to the key role predator risk effects can play in understanding conservation and wildlife management challenges. Here, we propose that population predator risk effects can be predicted through an extension of individual risk trade-off theory and show for the first time that this is the case in a wild vertebrate system. Specifically, we demonstrate that the timing (in specific months of the year), occurrence (at low food availability), cause (reduction in individual energy reserves), and type (starvation mortality) of a population-level predator risk effect can be successfully predicted from individual responses using a widely applicable theoretical framework (individual-based risk trade-off theory). Our results suggest that individual-based risk trade-off frameworks could allow a wide range of population-level predator risk effects to be predicted from existing ecological theory, which would enable risk effects to be more routinely integrated into consideration of population processes and in applied situations such as conservation.

Mass gained during breeding positively correlates with adult survival because both reflect life history adaptation to seasonal food availability.

Both mass (as a measure of body reserves) during breeding and adult survival should reflect variation in food availability. Those species that are adapted to less seasonally variable foraging niches and so where competition dominates during breeding, will tend to have a higher mass increase via an interrupted foraging response, because their foraging demands increase and so become more unpredictable. They will then produce few offspring per breeding attempt, but trade this off with higher adult survival. In contrast, those species that occupy a more seasonal niche will not gain mass because foraging remains predictable, as resources become superabundant during breeding. They can also produce more offspring per breeding attempt, but with a trade-off with reduced adult survival. We tested whether the then predicted positive correlation between levels of mass gained during seasonal breeding and adult survival was present across 40 species of tropical bird measured over a 10-year period in a West African savannah. We showed that species with a greater seasonal mass increase had higher adult survival, controlling for annual mass variation (i.e. annual variation in absolute food availability) and variation in the timing of peak mass (i.e. annual predictability of food availability), clutch size, body size, migratory status and phylogeny. Our results support the hypothesis that the degree of seasonal mass variation in birds is probably an indication of life history adaptation: across tropical bird species it may therefore be possible to use mass gain during breeding as an index of adult survival.

Predicting the optimal prey group size from predator hunting behaviour.

1. How group size affects predator attack and success rate, and so prey vulnerability, is important in determining the nonlethal consequences of predation risk on animal populations and communities. Theory predicts that both predator attack success rate and the dilution effect decline exponentially with group size and that selection generates optimal group sizes at a 'risk threshold' above which antipredation benefits are outweighed by costs, such as those owing to higher attack rates. 2. We examined whether flock size risk thresholds for attack rate, success rate or dilution differed, and therefore whether the strength of selection for group size differed for these three factors, using a system of redshank Tringa totanus flocks being hunted by Eurasian sparrowhawks Accipiter nisus. We also asked which of the three thresholds, on their own or in combination, predicted the most commonly observed group size. 3. Mean redshank flock size increased with a very gradual quadratic function (i.e. approximately linearly) with population size, although at a rate half that possible; when population size was not limiting, individuals almost always avoided flocks of less than 30 and birds were frequently in flocks up to at least 80. Sparrowhawk attack rate showed a quadratic relationship with flock size and peaked at 55 redshanks. Sparrowhawk attack success rate, however, declined exponentially, becoming less steep at flock sizes of about 40 and remaining uniformly low (a 95% decrease) by 70. Combined with dilution, individual risk of death per attack decreased by 95% when group size reached 30 (20 for the dilution effect alone). 4. Redshanks most commonly formed group sizes that gained the maximum individual predation risk reduction. They also commonly formed group sizes far above any further substantial advantages from the dilution effect or from reducing attack rate, but that continued to reduce predation risk by lowering attack success rate. Individuals did not always form the largest groups possible which we suggest is because individual variation in risk-taking subdivides the population. This places a constraint on the ability of individuals to compensate for predation risk and will have a variety of important effects on animal populations.