Accounting for imperfect detection when estimating species-area relationships and beta-diversity.

Ecologists have historically quantified fundamental biodiversity patterns, including species-area relationships (SARs) and beta diversity, using observed species counts. However, imperfect detection may often bias derived community metrics and subsequent community models. Although several statistical methods claim to correct for imperfect detection, their performance in species-area and ß-diversity research remains unproven. We examine inaccuracies in the estimation of SARs and ß-diversity parameters that emerge from imperfect detection, and whether such errors can be mitigated using a non-parametric diversity estimator (iNEXT.3D) and Multi-Species Occupancy Models (MSOMs). We simulated 28,350 sampling regimes of 2835 fragmented communities, varying the mean and standard deviation of species detection probabilities, and the number of sampling repetitions. We then quantified the bias, accuracy, and precision of derived estimates of model coefficients for SARs and the effects of patch area on ß-diversity (pairwise Sørensen similarity). Imperfect detection biased estimates of all evaluated parameters, particularly when mean detection probabilities were low, and there were few sampling repetitions. Observed counts consistently underestimated species richness and SAR z-values, and overestimated SAR c-values; iNEXT.3D and MSOMs only partially resolved these biases. iNEXT.3D provided the best estimates of SAR z-values, although MSOM estimates were generally comparable. All three methods accurately estimated pairwise Sørensen similarity in most circumstances, but only MSOMs provided unbiased estimates of the coefficients of models examining covariate effects on ß-diversity. Even when using iNEXT.3D or MSOMs, imperfect detection consistently caused biases in SAR coefficient estimates, calling into question the robustness of previous SAR studies. Furthermore, the inability of observed counts and iNEXT.3D to estimate ß-diversity model coefficients resulted from a systematic, area-related bias in Sørensen similarity estimates. Importantly, MSOMs corrected for these biases in ß-diversity assessments, even in suboptimal scenarios. Nonetheless, as estimator performance consistently improved with increasing sampling repetitions, the importance of appropriate sampling effort cannot be understated.

Movement of juvenile migratory birds from settlement to adulthood across the non-breeding range.

Among migratory vertebrates, high levels of fidelity to non-breeding sites during adulthood are common. If occupied sites vary in quality, strong site fidelity can have profound consequences for individual fitness and population demography. Given the prevalence of adult site fidelity, the regions of the non-breeding range to which juveniles first migrate, and the scale of any subsequent movements, are likely to be pivotal in shaping distributions and demographic processes across population ranges. However, inherent difficulties in tracking migratory individuals through early life mean that opportunities to quantify juvenile settlement and movements across non-breeding ranges, and the mechanisms involved, are extremely rare. Through long-term, range-wide resightings of hundreds of colour-marked individuals from their first migration to adulthood and the application of state-space models, we quantify levels of juvenile and adult regional-scale movements and distances at different life stages across the whole non-breeding distribution range in a migratory shorebird, the Black-tailed Godwit (Limosa limosa islandica). We show that the probability of individuals changing non-breeding regions (seven historical wintering regions spanning the Western Europe range) at all ages is very low (mean movement probability = 10.9% from first to subsequent winter, and 8.3% from first adult winter to later winters). Movement between regions was also low between autumn and winter of the same year for both juveniles (mean movement probability = 17.0%) and adults (10.4%). The great majority of non-breeding movements from the first autumn to adulthood were within regions and less than 100 km. The scarcity of regional-scale non-breeding movements from the first autumn to adulthood means that the factors influencing where juveniles settle will be key determinants of non-breeding distributions and of the rate and direction of changes in distributions.

Temperature and microclimate refugia use influence migratory timings of a threatened grassland bird.

BACKGROUND: Seasonal changes in resource availability are known to influence the migratory behaviour of animals, including both timing and distance. While the influence of environmental cues on migratory behaviour has been widely studied at the population level, it has rarely been examined at the spatial scale at which individuals experience their environment. Here, we test the hypothesis that individuals exposed to similar large-scale environmental cues may vary in migratory behaviour in response to the different microclimate conditions they experience at fine scales. METHODS: We combine high-spatial and temporal resolution microclimate and habitat information with GPS tracking data for a partially migratory threatened grassland bird. Data from 47 little bustards (Tetrax tetrax; 67 breeding events) tracked between 2009 and 2019 was used to (i) evaluate individual consistency in migratory behaviour (timing and distance) and (ii) assess whether the local environmental characteristics experienced by individuals - and in particular their use of microclimate refugia - influence distance and timing of migration, from and to the breeding sites. RESULTS: Migratory distance was consistent for birds tracked over multiple years, while the timing of migration showed high variability among individuals. Departures from breeding areas spanned from May to August, with a few birds remaining in their breeding areas. Vegetation greenness (a proxy for food availability) was positively associated with the time birds spent in the breeding area. The best model also included a positive effect of microclimate refugia availability on breeding season length, although an interaction with temperature suggested that this effect did not occur at the highest relative temperatures. The return date to breeding grounds, although spanning from September to April, was not influenced by the environmental conditions or food availability. CONCLUSIONS: Food availability, measured by a vegetation greenness proxy, was associated with later migration at the end of the breeding season. Availability of cooler microclimate refugia may also allow for later departures from the breeding sites in all but the hottest conditions. Management measures that increase microclimate refugia availability and provide foraging resources can thus potentially increase the length of the breeding season for this species.

Combining remote sensing and tracking data to quantify species' cumulative exposure to anthropogenic change.

Identifying when and where organisms are exposed to anthropogenic change is crucial for diagnosing the drivers of biodiversity declines and implementing effective conservation measures. Accurately measuring individual-scale exposure to anthropogenic impacts across the annual cycle as they move across continents requires an approach that is both spatially and temporally explicit-now achievable through recent parallel advances in remote-sensing and individual tracking technologies. We combined 10 years of tracking data for a long-distance migrant, (common cuckoo, Cuculus canorus), with multi-dimensional remote-sensed spatial datasets encompassing thirteen relevant anthropogenic impacts (including infrastructure, hunting, habitat change, and climate change), to quantify mean hourly and total accumulated exposure of tracked individuals to anthropogenic change across each stage of the annual cycle. Although mean hourly exposure to anthropogenic change was greatest in the breeding stage, accumulated exposure to changes associated with direct mortality risks (e.g., built infrastructure) and with climate were greatest during the wintering stage, which comprised 63% of the annual cycle on average for tracked individuals. Exposure to anthropogenic change varied considerably within and between migratory flyways, but there were no clear between-flyway differences in overall exposure during migration stages. However, more easterly autumn migratory routes were significantly associated with lower subsequent exposure to anthropogenic impacts in the winter stage. Cumulative change exposure was not significantly associated with recent local-scale population trends in the breeding range, possibly because cuckoos from shared breeding areas may follow divergent migration routes and therefore encounter very different risk landscapes. Our study highlights the potential for the integration of tracking data and high-resolution remote sensing to generate valuable and detailed new insights into the impacts of environmental change on wild species.

Combining bird tracking data with high-resolution thermal mapping to identify microclimate refugia.

Elevated temperatures can have a range of fitness impacts, including high metabolic cost of thermoregulation, hence access to microclimate refugia may buffer individuals against exposure to high temperatures. However, studies examining the use of microclimate refugia, remain scarce. We combined high resolution microclimate modelling with GPS tracking data as a novel approach to identify the use and availability of cooler microclimate refugia (sites > 0.5 °C cooler than the surrounding landscape) at the scales experienced by individual animals. 77 little bustards (Tetrax tetrax) were tracked between 2009 and 2019. The 92,685 GPS locations obtained and their surrounding 500 m areas were characterised with hourly temperature and habitat information at 30 m × 30 m and used to determine microclimate refugia availability and use. We found that the semi-natural grassland landscapes used by little bustards have limited availability of cooler microclimate areas-fewer than 30% of the locations. The use of cooler microclimate sites by little bustards increased at higher ambient temperatures, suggesting that individuals actively utilise microclimate refugia in extreme heat conditions. Microclimate refugia availability and use were greater in areas with heterogeneous vegetation cover, and in coastal areas. This study identified the landscape characteristics that provide microclimate opportunities and shelter from extreme heat conditions. Little bustards made greater use of microclimate refugia with increasing temperatures, particularly during the breeding season, when individuals are highly site faithful. This information can help identify areas where populations might be particularly exposed to climate extremes due to a lack of microclimate refugia, and which habitat management measures may buffer populations from expected increased exposure to temperature extremes.

Biogeographic multi-species occupancy models for large-scale survey data.

Ecologists often seek to infer patterns of species occurrence or community structure from survey data. Hierarchical models, including multi-species occupancy models (MSOMs), can improve inference by pooling information across multiple species via random effects. Originally developed for local-scale survey data, MSOMs are increasingly applied to larger spatial scales that transcend major abiotic gradients and dispersal barriers. At biogeographic scales, the benefits of partial pooling in MSOMs trade off against the difficulty of incorporating sufficiently complex spatial effects to account for biogeographic variation in occupancy across multiple species simultaneously. We show how this challenge can be overcome by incorporating preexisting range information into MSOMs, yielding a "biogeographic multi-species occupancy model" (bMSOM). We illustrate the bMSOM using two published datasets: Parulid warblers in the United States Breeding Bird Survey and entire avian communities in forests and pastures of Colombia's West Andes. Compared with traditional MSOMs, the bMSOM provides dramatically better predictive performance at lower computational cost. The bMSOM avoids severe spatial biases in predictions of the traditional MSOM and provides principled species-specific inference even for never-observed species. Incorporating preexisting range data enables principled partial pooling of information across species in large-scale MSOMs. Our biogeographic framework for multi-species modeling should be broadly applicable in hierarchical models that predict species occurrences, whether or not false absences are modeled in an occupancy framework.

Bird migration: When vagrants become pioneers.

Vagrant birds are frequently recorded outside of their regular geographic range. A new study documents how vagrancy, in this case in an Asian songbird, can lead to establishment of a new migration route.

Sensitivity of migratory connectivity estimates to spatial sampling design.

BACKGROUND: The use of statistical methods to quantify the strength of migratory connectivity is commonplace. However, little attention has been given to their sensitivity to spatial sampling designs and scales of inference. METHODS: We examine sources of bias and imprecision in the most widely used methodology, Mantel correlations, under a range of plausible sampling regimes using simulated migratory populations. RESULTS: As Mantel correlations depend fundamentally on the spatial scale and configuration of sampling, unbiased inferences about population-scale connectivity can only be made under certain sampling regimes. Within a contiguous population, samples drawn from smaller spatial subsets of the range generate lower connectivity metrics than samples drawn from the range as a whole, even when the underlying migratory ecology of the population is constant across the population. Random sampling of individuals from contiguous subsets of species ranges can therefore underestimate population-scale connectivity. Where multiple discrete sampling sites are used, by contrast, overestimation of connectivity can arise due to samples being biased towards larger between-individual pairwise distances in the seasonal range where sampling occurs (typically breeding). Severity of all biases was greater for populations with lower levels of true connectivity. When plausible sampling regimes were applied to realistic simulated populations, accuracy of connectivity measures was maximised by increasing the number of discrete sampling sites and ensuring an even spread of sites across the full range. CONCLUSIONS: These results suggest strong potential for bias and imprecision when making quantitative inferences about migratory connectivity using Mantel statistics. Researchers wishing to apply these methods should limit inference to the spatial extent of their sampling, maximise their number of sampling sites, and avoid drawing strong conclusions based on small sample sizes.

Replacing low-intensity cattle pasture with oil palm conserves dung beetle functional diversity when paired with forest protection.

Meeting rising demand for oil palm whilst minimizing the loss of tropical biodiversity and associated ecosystem functions is a core conservation challenge. One potential solution is focusing the expansion of high-yielding crops on presently low-yielding farmlands alongside protecting nearby tropical forests that can enhance provision of ecosystem functions. A key question is how this solution would impact invertebrate functional diversity. We focus on oil palm in the Colombian Llanos, where plantations are replacing improved cattle pastures and forest fragments, and on dung beetles, which play key functional roles in nutrient cycling and secondary seed dispersal. We show that functional richness and functional diversity of dung beetles is greater in oil palm than in cattle pasture, and that functional metrics did not differ between oil palm and remnant forest. The abundance-size class profile of dung beetles in oil palm was more similar to forest than to pasture, which had lower abundances of the smallest and largest dung beetles. The abundance of tunneling and rolling dung beetles did not differ between oil palm and forest, while higher forest cover increased the abundance of diurnal and generalist-feeding beetles in oil palm landscapes. This suggests that prioritizing agricultural development on low-yielding cattle pasture will have positive effects on functional diversity and highlights the need for forest protection to maintain ecosystem functioning within agricultural landscapes.

Sparing land for secondary forest regeneration protects more tropical biodiversity than land sharing in cattle farming landscapes.

Effectively managing farming to meet food demand is vital for the future of biodiversity.1,2 Increasing yields on existing farmland can allow the abandonment (sparing) of low-yielding areas that subsequently recover as secondary forest.2-5 A key question is whether such "secondary sparing" conserves biodiversity more effectively than retaining wildlife-friendly habitat within farmland ("land sharing"). Focusing on the Colombian Choco-Andes, a global hotspot of threatened biodiversity,6 and on cattle farming, we examined the outcomes of secondary sparing and land sharing via simulated scenarios that maintained constant landscape-wide production and equal within-pasture yield: (1) for species and functional diversity of dung beetles and birds; (2) for avian phylogenetic diversity; and (3) across different stages of secondary forest regeneration, relative to spared primary forests. Sparing older secondary forests (15-30 years recovery) promotes substantial species, functional, and phylogenetic (birds only) diversity benefits for birds and dung beetles compared to land sharing. Species of conservation concern had higher occupancy estimates under land-sparing compared to land-sharing scenarios. Spared secondary forests accumulated equivalent diversity to primary forests for dung beetles within 15 years and within 15-30 years for birds, highlighting the need for longer term protection to maximize the biodiversity gains of secondary sparing. Promoting the recovery and protection of large expanses of secondary forests under the land-sparing model provides a critical mechanism for protecting tropical biodiversity, with important implications for concurrently assisting in the delivery of global targets to restore 350 million hectares of forested landscapes.7,8.

Carryover effects of long-distance avian migration are weaker than effects of breeding environment in a partially migratory bird.

Migration may expose individuals to a wide range of increasing anthropogenic threats. In addition to direct mortality effects, this exposure may influence post-migratory reproductive fitness. Partial migration-where a population comprises migrants and residents-represents a powerful opportunity to explore carryover effects of migration. Studies of partial migration in birds typically examine short-distance systems; here we studied an unusual system where residents breed in mixed colonies alongside long-distance trans-Saharan migrants (lesser kestrels (Falco naumanni) in Spain). Combining geolocator data, stable isotope analysis and resighting data, we examined the effects of this stark difference in migratory strategy on body condition, breeding phenology and breeding success. We monitored four colonies in two regions of southern Spain for five consecutive years (2014-2018), yielding 1962 captures, determining migratory strategy for 141 adult bird-years. Despite a 3000-km difference in distance travelled, we find no effect of strategy on breeding parameters. We find weak evidence for a short-term negative carryover effect of migration on body condition, but this was only apparent in the breeding region with lower primary productivity. Our results indicate that carryover effects of even highly divergent migratory strategies may be minimal relative to effects of conditions experienced on breeding grounds.

Mass-abundance scaling in avian communities is maintained after tropical selective logging.

Selective logging dominates forested landscapes across the tropics. Despite the structural damage incurred, selectively logged forests typically retain more biodiversity than other forest disturbances. Most logging impact studies consider conventional metrics, like species richness, but these can conceal subtle biodiversity impacts. The mass-abundance relationship is an integral feature of ecological communities, describing the negative relationship between body mass and population abundance, where, in a system without anthropogenic influence, larger species are less abundant due to higher energy requirements. Changes in this relationship can indicate community structure and function changes.We investigated the impacts of selective logging on the mass-abundance scaling of avian communities by conducting a meta-analysis to examine its pantropical trend. We divide our analysis between studies using mist netting, sampling the understory avian community, and point counts, sampling the entire community.Across 19 mist-netting studies, we found no consistent effects of selective logging on mass-abundance scaling relative to primary forests, except for the omnivore guild where there were fewer larger-bodied species after logging. In eleven point-count studies, we found a more negative relationship in the whole community after logging, likely driven by the frugivore guild, showing a similar pattern.Limited effects of logging on mass-abundance scaling may suggest high species turnover in logged communities, with like-for-like replacement of lost species with similar-sized species. The increased negative mass-abundance relationship found in some logged communities could result from resource depletion, density compensation, or increased hunting; potentially indicating downstream impacts on ecosystem functions. Synthesis and applications. Our results suggest that size distributions of avian communities in logged forests are relatively robust to disturbance, potentially maintaining ecosystem processes in these forests, thus underscoring the high conservation value of logged tropical forests, indicating an urgent need to focus on their protection from further degradation and deforestation.

Fitness consequences of different migratory strategies in partially migratory populations: A multi-taxa meta-analysis.

Partial migration-wherein migratory and non-migratory individuals exist within the same population-represents a behavioural dimorphism; for it to persist over time, both strategies should yield equal individual fitness. This balance may be maintained through trade-offs where migrants gain survival benefits by avoiding unfavourable conditions, while residents gain breeding benefits from early access to resources. There has been little overarching quantitative analysis of the evidence for this fitness balance. As migrants-especially long-distance migrants-may be particularly vulnerable to environmental change, it is possible that recent anthropogenic impacts could drive shifts in fitness balances within these populations. We tested these predictions using a multi-taxa meta-analysis. Of 2,939 reviewed studies, 23 contained suitable information for meta-analysis, yielding 129 effect sizes. Of these, 73% (n = 94) reported higher resident fitness, 22% (n = 28) reported higher migrant fitness, and 5% (n = 7) reported equal fitness. Once weighted for precision, we found balanced fitness benefits across the entire dataset, but a consistently higher fitness of residents over migrants in birds and herpetofauna (the best-sampled groups). Residency benefits were generally associated with survival, not breeding success, and increased with the number of years of data over which effect sizes were calculated, suggesting deviations from fitness parity are not due to sampling artefacts. A pervasive survival benefit to residency documented in recent literature could indicate that increased exposure to threats associated with anthropogenic change faced by migrating individuals may be shifting the relative fitness balance between strategies.

La migration partielle - dans laquelle les migrants et les résidents existent dans une seul population - représente un dimorphisme comportemental; pour qu'elles persiste au fil du temps, les deux stratégies doivent conférer la même valeur adaptive (fitness) individuelle. Cet équilibre peut être maintenu par un compromis où les migrants obtiennent des bénéfices de survie en évitant des conditions défavorables, tandis que les résidents obtiennent des avantages reproductifs grâce á un accès précoce aux ressources. Peu d'analyses quantitatives ont été entreprise sur les preuves de cet équilibre de fitness. Étant donné que les migrants - en particulier les migrants de longue distance - peuvent être particulièrement vulnérables aux changements environnementaux, il est possible que les impacts anthropiques récents puissent entraîner une modification dans l'équilibre de fitness au sein de ces populations. Nous avons testé ces prédictions en utilisant une méta-analyse multi-taxa. Sur 2,939 études examinées, 23 contenaient des informations appropriées pour la méta-analyse, donnant 129 tailles d'effet. Parmi eux, 73% (n = 94) ont déclaré un fitness supérieur pour les résidents, 22% (n = 28) un fitness supérieur pour les migrants, et 5% (n = 7) une égalité de fitness entre les deux. Après avoir pondéré les données par la précision, nous avons trouvé des avantages équilibrés sur le fitness sur l'ensemble des données, mais parmi les oiseaux et l'herpétofaune (les groupes les mieux échantillonnés), de manière cohérente, les résidents étaient associés à un meilleur fitness comparé à celle des migrants. Les bénéfices d'être résident étaient associés à la survie, et non au succès reproductif, et augmente avec le nombre d'années de données pendant lesquelles la taille d'effet a été calculé; cela suggérerait que les écarts par rapport à la parité de fitness ne sont pas dus à des artefacts d'échantillonnage. Cet avantage de survie pour les résidents, documenté dans la littérature récente, pourrait indiquer qu'une exposition accrue aux menaces associées aux changements anthropiques rencontrés par les individus migrateurs pourrait modifier l'équilibre entre les stratégies.

The impact of secondary forest regeneration on ground-dwelling ant communities in the Tropical Andes.

Natural regeneration of abandoned farmland provides an important opportunity to contribute to global reforestation targets, including the Bonn Challenge. Of particular importance are the montane tropics, where a long history of farming, frequently on marginal soils, has rendered many ecosystems highly degraded and hotspots of extinction risk. Ants play crucial roles in ecosystem functioning, and a key question is how time since abandonment and elevation (and inherent temperature gradients therein) affect patterns of ant recovery within secondary forest systems. Focusing on the Colombian Andes across a 1300 m altitudinal gradient and secondary forest (2-30 years) recovering on abandoned cattle pastures, we find that over time ant community composition and species richness recovered towards that of primary forest. However, these relationships are strongly dependent on elevation with the more open and warmer pasturelands supporting more ants than either primary or secondary forest at a particular elevation. The loss of species richness and change in species composition with elevation is less severe in pasture than forests, suggesting that conditions within pasture and its remaining scattered trees, hedgerows and forest fragments, are more favourable for some species, which are likely in or near thermal debt. Promoting and protecting natural regenerating forests over the long term in the montane tropics will likely offer significant potential for returning ant communities towards primary forest levels.

Land-sparing agriculture sustains higher levels of avian functional diversity than land sharing.

The ecological impacts of meeting rising demands for food production can potentially be mitigated by two competing land-use strategies: off-setting natural habitats through intensification of existing farmland (land sparing), or elevating biodiversity within the agricultural matrix via the integration of "wildlife-friendly" habitat features (land sharing). However, a key unanswered question is whether sparing or sharing farming would best conserve functional diversity, which can promote ecosystem stability and resilience to future land-use change. Focusing on bird communities in tropical cloud forests of the Colombian Andes, we test the performance of each strategy in conserving functional diversity. We show that multiple components of avian functional diversity in farmland are positively related to the proximity and extent of natural forest. Using landscape and community simulations, we also show that land-sparing agriculture conserves greater functional diversity and predicts higher abundance of species supplying key ecological functions than land sharing, with sharing becoming progressively inferior with increasing isolation from remnant forest. These results suggest low-intensity agriculture is likely to conserve little functional diversity unless large blocks of adjacent natural habitat are protected, consistent with land sparing. To ensure the retention of functionally diverse ecosystems, we urgently need to implement mechanisms for increasing farmland productivity whilst protecting spared land.

Impacts of tropical forest disturbance on species vital rates.

Tropical forests are experiencing enormous threats from deforestation and habitat degradation. Much knowledge of the impacts of these land-use changes on tropical species comes from studies examining patterns of richness and abundance. Demographic vital rates (survival, reproduction, and movement) can also be affected by land-use change in a way that increases species vulnerability to extirpation, but in many cases these impacts may not be manifested in short-term changes in abundance or species richness. We conducted a literature review to assess current knowledge and research effort concerning how land-use change affects species vital rates in tropical forest vertebrates. We found a general paucity of empirical research on demography across taxa and regions, with some biases toward mammals and birds and land-use transitions, including fragmentation and agriculture. There is also considerable between-species variation in demographic responses to land-use change, which could reflect trait-based differences in species sensitivity, complex context dependencies (e.g., between-region variation), or inconsistency in methods used in studies. Efforts to improve understanding of anthropogenic impacts on species demography are underway, but there is a need for increased research effort to fill knowledge gaps in understudied tropical regions and taxa. The lack of information on demographic impacts of anthropogenic disturbance makes it difficult to draw definite conclusions about the magnitude of threats to tropical ecosystems under anthropogenic pressures. Thus, determining conservation priorities and improving conservation effectiveness remains a challenge.

Interspecific variation in responses to microclimate by terrestrial isopods: implications in relation to climate change.

The importance of considering species-specific biotic interactions when predicting feedbacks between the effects of climate change and ecosystem functions is becoming widely recognised. The responses of soil animals to predicted changes in global climate could potentially have far-reaching consequences for fluxes of soil carbon, including climatic feedbacks resulting from increased emissions of carbon dioxide from soils. The responses of soil animals to different microclimates can be summarised as norms of reaction, in order to compare phenotypic differences in traits along environmental gradients. Thermal and moisture reaction norms for physiological, behavioural and life history traits of species of terrestrial isopods differing in their morphological adaptations for reducing water loss are presented. Gradients of moisture reaction norms for respiratory rates and thermal reaction norms for water loss, for a species from the littoral zone were steeper than those for species from mesic environments. Those for mesic species were steeper than for those from xeric habitats. Within mesic species, gradients of thermal reaction norms for aggregation were steeper for Oniscusasellus than for Porcellioscaber or Armadilliumvulgare, and moisture reaction norms for sheltering and feeding behaviours were steeper for Philosciamuscorum than for either P.scaber or A.vulgare. These differences reflect differences in body shape, permeability of the cuticle, and development of pleopodal lungs. The implications of differences between different species of soil animals in response to microclimate on the possible influence of the soil fauna on soil carbon dynamics under future climates are discussed. In conclusion a modelling approach to bridging the inter-disciplinary gap between carbon cycling and the biology of soil animals is recommended.

Stay-at-home strategy brings fitness benefits to migrants.

In Focus: Lok, T., Veldhoen, L., Overdijk, O., Tinbergen, J. M., & Piersma, T. (2017). An age-dependent fitness cost of migration? Old trans-Saharan migrating spoonbills breed later than those staying in Europe, and late breeders have lower recruitment. Journal of Animal Ecology, 86, 998-1009. In Focus: Grist, H., Daunt, F., Wanless, S., Burthe, S. J., Newell, M. A., Harris, M. P., & Reid, J. M. (2017). Reproductive performance of resident and migrant males, females and pairs in a partially migratory bird. Journal of Animal Ecology, 86, 1010-1021. In this issue, two studies examine the extent to which variation in migratory behaviour influences individual fitness across a population. Lok, Veldhoen, Overdijk, Tinbergen, and Piersma () examine reproductive success and post-fledging survival in a population of Eurasian spoonbills (Platalea leucorodia), comparing individuals that winter in south-west Europe against those migrating to sub-Saharan Africa, while Grist et al. () measure reproductive success in a population of European shags (Phalacrocorax aristotelis) breeding in Scotland that either remain resident or migrate to surrounding waters. Both studies find that individuals migrating longer distances tend to show later initiation of breeding attempts. In turn, longer migration correlates with lower reproductive success in both populations. In spoonbills, this effect is most pronounced in older male birds, while young individuals show little difference in breeding success with respect to migration distance. In shags, fitness benefits of residence were most pronounced when both individuals of a pair were resident, although there was no evidence of assortative mating. Both studies provide fascinating new insights into the role migratory variability can play in shaping population dynamics.

Simple settlement decisions explain common dispersal patterns in territorial species.

Dispersal is one of the least-understood aspects of animal behaviour. For example, little is known of the mechanisms that determine how individuals express different dispersal behaviours in response to different circumstances. Uncovering these mechanisms is important for our understanding of spatial population dynamics. Using agent-based simulations, we examine how simple decision rules generate individual-level dispersal plasticity, and how this can influence population-scale dispersal dynamics. We model a territorial, monogamous population inhabiting a completely homogeneous environment. Dispersal variability therefore emerges solely as a result of between-individual interactions (competition, settlement, reproduction), which are governed by simple decision-making algorithms. We show that complex dispersal dynamics, including sex biases and strong density dependence, emerge naturally from simple rule-based behaviours. Dispersal is particularly sensitive to the inclusion of mate availability as a criterion for settlement: if neither sex evaluates mate availability, dispersal distances tend to decline at low densities, leading to a strong Allee effect from reduced pairing success. If one sex evaluates mate availability (females), Allee effects are largely avoided, but female-biased dispersal generates increasingly male-biased adult sex ratios at low densities. Sex biases are eliminated if both sexes evaluate mate availability, but population growth rates tend to be reduced due to survival costs and reduced pairing success. Our models suggest that simple decision mechanisms can explain several dynamic patterns that are commonly observed among territorial species. Importantly, these patterns emerge in the absence of environmental heterogeneity or between-individual variation in dispersal phenotypes, two conditions that are often invoked to explain dispersal heterogeneity in nature. This has implications for studies seeking to examine the causes of dispersal variability in wild populations, suggesting that observed patterns could be largely driven by the social and demographic conditions experienced by sampled individuals. Further insights could be gained by examining how selection operates on decision rules in different life-history and environmental circumstances, and how this might interact with selection on other demographic traits. Uncovering the decision rules used during settlement should be a priority for those wishing to understand and predict dispersal patterns in nature.