Mortality-minimizing sandpipers vary stopover behavior dependent on age and geographic proximity to migrating predators

Ecological theory for long-distance avian migration considers time-, energy-, and mortality-minimizing tactics, but predictions about the latter have proven elusive. Migrants must make behavioral decisions that can favor either migratory speed or safety from predators, but often not both. We compare the behavior of adult and juvenile western sandpipers Calidris mauri during the course of their temporally segregated passages at a major stopover site. Here, the passage and winter arrival of an important predator, the peregrine falcon Falco peregrinus begins near the end of the adult sandpiper passage (July) and increases rapidly through the juvenile passage (August). The mortality-minimizing hypothesis predicts that as the falcon front is distant but approaching, sandpipers should initially increase the fuel-loading rate (lowered vigilance and predator apprehension) to increase migration speed and so maintain their head start. As the falcon front gains proximity to and passes over the stopover site, sandpipers should become increasingly cautious. Our measurements show that adults decreased vigilance during the period prior to falcon arrival, and had lower vigilance overall than juveniles. Juveniles were more apprehensive, flying further and longer in response to disturbance by a falcon silhouette. This trend was reversed in response to a human approach. Both groups were more vigilant and more apprehensive in a study year with earlier falcon arrival. These results suggest that late (juvenile) and early (adult) migrants minimize mortality on migration in different ways, adults by increased migratory speed at the expense of caution on stopover sites, and juveniles by increased caution at the expense of speed.     

A method for the detection of non-random associations among flocking birds and its application to sanderlings Calidris alba wintering in N.E. England

Summary Indices of association are frequently used in studies of social behaviour. Observed association indices may be compared with those obtained from computer randomizations of the data in order to test whether any two individuals in a population tend to occur together more than would be expected by chance. Data for individually-identifiable colour-ringed sanderlings Calidris alba are presented to illustrate the importance of the assumptions on which such null models are built. When each individual was given an equal chance of appearing in each flock associations differed from those expected by chance. But when the different numbers of times that each individual occurred at the study site were taken into account associations were not different from those expected by chance. Hypotheses relating association levels to a number of factors were tested. None of the factors examined had significant and appreciable effects on association levels. In particular, there was no evidence for sanderlings moving around in stable groups. Taking into account the different distributions of different individuals, the population of birds present on any day represented a random sample from the population of birds seen during the winter. Correspondence to: G. Roberts     

The Impact of Variation in Stopover Habitat Quality on Migrant Rufous Hummingbirds

We examined the impact of variation in habitat quality on migrating Rufous Hummingbirds ( Selasphorus rufus ) in the California Sierra Nevada. As do other migratory species, these birds depend on "stopover" habitats en route for feeding and replenishing depleted energy stores. During seven years of study, the quality of the stopover habitat (assessed in terms of the density of nectar food resources) varied widely due to natural variation in flowering. In years when stopover habitat quality was poor, incoming body masses were low and stopover durations were long. Population densities of migrant hummingbirds at the study site were coupled to habitat quality both within and among years. These observations demonstrate important effects of stopover habitat variation on the physiological, behavioral, and population ecology of migrating hummingbirds. High-quality stopover habitats are critical links between breeding and wintering areas for many species, and their preservation should be considered an essential component of strategies aiming to conserve migratory bird populations.     

A field investigation of scrounging in semipalmated sandpipers

Animals that forage in groups can produce their own food patches or scrounge the food discoveries of their companions. Mean tactic payoffs are expected to be the same at equilibrium for phenotypically equal foragers. Scrounging is also typically viewed as a risk-averse foraging strategy that provides a more even food intake rate over time. The occurrence of scrounging and the payoffs from different foraging modes have rarely been investigated in the field. Over two field seasons, I examined patch sharing in semipalmated sandpipers (Calidris pusilla) foraging on minute food items at the surface of the substrate. Birds could find patches on their own, a producing event, or join the food patches discovered by others, a scrounging event. I found that the average search time per patch did not differ between producing and scrounging but that the average time spent exploiting a patch was reduced nearly by half when scrounging. As a result, the proportion of time spent exploiting a patch, a measure of foraging payoffs, was significantly lower when scrounging. The variance in payoffs was similar for producing and scrounging. When producing their own patches, individuals that scrounged spent the same proportion of time exploiting a patch as those that only produced. However, within the same individuals, the search time for a scrounged patch was longer than the search time for a produced patch. The results show unequal payoffs for producing and scrounging in this system and suggest that low success in finding patches elicited scrounging.     

Effect of gyrodactylid ectoparasites on host behaviour and social network structure in guppies <Emphasis Type="Italic">Poecilia reticulata</Emphasis>

Understanding how individuals modify their social interactions in response to infectious disease is of central importance for our comprehension of how disease dynamics operate in real-world populations. Whilst a significant amount of theoretical work has modelled disease transmission using network models, we have comparatively little understanding of how infectious disease impacts on the social behaviour of individuals and how these effects scale up to the level of the population. We experimentally manipulated the parasite load of female guppies (Poecilia reticulata) and introduced fish either infected with the ectoparasites Gyrodactylus spp. (experimental) or uninfected (control) into replicated semi-natural populations of eight size-matched female guppies. We quantified the behaviour and social associations of both the introduced fish and the population fish. We found that infected experimental fish spent less time associating with the population fish than the uninfected control fish. Using information on which fish initiated shoal fission (splitting) events, our results demonstrate that the population fish actively avoided infected experimental fish. We also found that the presence of an infected individual resulted in a continued decline in social network clustering up to at least 24 h after the introduction of the infected fish, whereas in the control treatment, the clustering coefficient showed an increase at this time point. These results demonstrate that the presence of a disease has implications for both the social associations of infected individuals and for the social network structure of the population, which we predict will have consequences for infectious disease transmission.     

Characterizing source-sink dynamics with genetic parentage assignments

Source-sink dynamics have been suggested to characterize the population structure of many species, but the prevalence of source-sink systems in nature is uncertain because of inherent challenges in estimating migration rates among populations. Migration rates are often difficult to estimate directly with demographic methods, and indirect genetic methods are subject to a variety of assumptions that are difficult to meet or to apply to evolutionary timescales. Furthermore, such methods cannot be rigorously applied to high-gene-flow species. Here, we employ genetic parentage assignments in conjunction with demographic simulations to infer the level of immigration into a putative sink population. We use individual-based demographic models to estimate expected distributions of parent-offspring dyads under competing sink and closed-population models. By comparing the actual number of parent-offspring dyads (identified from multilocus genetic profiles) in a random sample of individuals taken from a population to expectations under these two contrasting demographic models, it is possible to estimate the rate of immigration and test hypotheses related to the role of immigration on population processes on an ecological timescale. The difference in the expected number of parent-offspring dyads between the two population models was greatest when immigration into the sink population was high, indicating that unlike traditional population genetic inference models, the highest degree of statistical power is achieved for the approach presented here when migration rates are high. We used the proposed genetic parentage approach to demonstrate that a threatened population of Marbled Murrelets (Braclhyrarmphus marmotus) appears to be supplemented by a low level of immigration (approximately 2-6% annually) from other populations.     

Prospecting at conspecific nests and exploration in a novel environment are associated with reproductive success in the jackdaw

Consistent behavioural differences between individuals of the same population (“personality” variation) might arise if individuals follow different life-history strategies. Thus, it would be important to determine how personality variation relates to behaviours potentially associated with life-history strategies, such as those related to the use of information about the state of the environment. Little is, however, known about how personality is associated with information use and reproductive success. We tested whether wild social jackdaws, Corvus monedula, show consistent behavioural differences in their exploratory behaviour (in a novel environment in the lab and in their reaction towards a novel object in the wild) and prospecting behaviour (number of visits to conspecific nests). We furthermore examined whether these behavioural traits are linked with each other and predictors of reproductive success. Breeding jackdaws were consistent in their exploratory behaviour within, and in their prospecting behaviour between, years. Exploratory behaviour in the novel environment was correlated with the latency to approach a novel object in the wild but not with the frequency of prospecting at conspecific nests. Highly exploratory males and females and frequently prospecting males produced fewer fledglings than less exploratory individuals or less prospecting males, respectively. We discuss the importance of consistent individual differences in exploration and information sampling on individual fitness.     

Simultaneous-count models to estimate abundance from counts of unmarked individuals with imperfect detection

We developed a method to estimate population abundance from simultaneous counts of unmarked individuals over multiple sites. We considered that at each sampling occasion, individuals in a population could be detected at 1 of the survey sites or remain undetected and used either multinomial or binomial simultaneous-count models to estimate abundance, the latter being equivalent to an N-mixture model with one site. We tested model performance with simulations over a range of detection probabilities, population sizes, growth rates, number of years, sampling occasions, and sites. We then applied our method to 3 critically endangered vulture species in Cambodia to demonstrate the real-world applicability of the model and to provide the first abundance estimates for these species in Cambodia. Our new approach works best when existing methods are expected to perform poorly (i.e., few sites and large variation in abundance among sites) and if individuals may move among sites between sampling occasions. The approach performed better when there were >8 sampling occasions and net probability of detection was high (>0.5). We believe our approach will be useful in particular for simultaneous surveys at aggregation sites, such as roosts. The method complements existing approaches for estimating abundance of unmarked individuals and is the first method designed specifically for simultaneous counts.     

Reserve design to optimize functional connectivity and animal density

Ecological distance-based spatial capture–recapture models (SCR) are a promising approach for simultaneously estimating animal density and connectivity, both of which affect spatial population processes and ultimately species persistence. We explored how SCR models can be integrated into reserve-design frameworks that explicitly acknowledge both the spatial distribution of individuals and their space use resulting from landscape structure. We formulated the design of wildlife reserves as a budget-constrained optimization problem and conducted a simulation to explore 3 different SCR-informed optimization objectives that prioritized different conservation goals by maximizing the number of protected individuals, reserve connectivity, and density-weighted connectivity. We also studied the effect on our 3 objectives of enforcing that the space-use requirements of individuals be met by the reserve for individuals to be considered conserved (referred to as home-range constraints). Maximizing local population density resulted in fragmented reserves that would likely not aid long-term population persistence, and maximizing the connectivity objective yielded reserves that protected the fewest individuals. However, maximizing density-weighted connectivity or preemptively imposing home-range constraints on reserve design yielded reserves of largely spatially compact sets of parcels covering high-density areas in the landscape with high functional connectivity between them. Our results quantify the extent to which reserve design is constrained by individual home-range requirements and highlight that accounting for individual space use in the objective and constraints can help in the design of reserves that balance abundance and connectivity in a biologically relevant manner.     

Predator inspection behaviour in three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>): body size,local predation pressure and cooperation

The apparently maladaptive tendency of fish to approach and inspect potential predators has been explained in terms of useful information gathering or as a signal to the predator that it has been seen. We examined this behaviour in 16 populations of wild-caught stickleback (Gasterosteus aculeatus) from ponds with and without predatory perch (Perca fluviatilis). Three large and three small individuals per population were each exposed to three model predators differing in realism. A final cooperative treatment entailed pairing subjects with a second individual from the same population, but of the alternative size class, during predator presentation. As might be expected, predator inspection behaviour was much greater in the predator-sympatric populations, and only these fish increased their level of inspection as the models became incrementally more realistic. This suggests that reductions occur in the level of costly inspection behaviour in populations without predators. Subject body size had no effect on inspection effort, which suggests a limited role for experience (we assumed larger fish to be older than smaller fish), at least over the relative age differences utilized. However, small predator-sympatric fish were the only subjects to increase inspection significantly when in a cooperative context, perhaps reflecting the inherent value of a relatively larger partner in this context. These results confirm that levels of predator inspection are both population- and situation-dependent, suggesting a trade-off in the potential costs and benefits of this behaviour.Communicated by C. St. Mary     

Predicting and Mapping Potential Whooping Crane Stopover Habitat to Guide Site Selection for Wind Energy Projects

Migratory stopover habitats are often not part of planning for conservation or new development projects. We identified potential stopover habitats within an avian migratory flyway and demonstrated how this information can guide the site‐selection process for new development. We used the random forests modeling approach to map the distribution of predicted stopover habitat for the Whooping Crane (Grus americana), an endangered species whose migratory flyway overlaps with an area where wind energy development is expected to become increasingly important. We then used this information to identify areas for potential wind power development in a U.S. state within the flyway (Nebraska) that minimize conflicts between Whooping Crane stopover habitat and the development of clean, renewable energy sources. Up to 54% of our study area was predicted to be unsuitable as Whooping Crane stopover habitat and could be considered relatively low risk for conflicts between Whooping Cranes and wind energy development. We suggest that this type of analysis be incorporated into the habitat conservation planning process in areas where incidental take permits are being considered for Whooping Cranes or other species of concern. Field surveys should always be conducted prior to construction to verify model predictions and understand baseline conditions. Predicción y Mapeo del Hábitat Potencial de Descanso de la Grulla Americana para Guiar la Selección de Sitios para Proyectos de Energía Eólica.     

A general random walk model for the leptokurtic distribution of organism movement: Theory and application

The movement of organisms is usually leptokurtic in which some individuals move long distances while the majority remains at or near the area they are released. There has been extensive research into the origin of such leptokurtic movement, but one important aspect that has been overlooked is that the foraging behaviour of most organisms is not Brownian as assumed in most existing models. In this paper we show that such non-Brownian foraging indeed gives rise to leptokurtic distribution. We first present a general random walk model to describe the organism movement by breaking the foraging of each individual into events of active movement and inactive stationary period; its foraging behaviour is therefore fully characterized by a joint probability of how far the individual can move in each active movement and the duration it remains stationary between two consecutive movements. The spatio-temporal distribution of the organism can be described by a generalized partial differential equation, and the leptokurtic distribution is a special case when the stationary period is not exponentially distributed. Empirical observations of some organisms living in different habitats indicated that their rest time shows a power-law distribution, and we speculate that this is general for other organisms. This leads to a fractional diffusion equation with three parameters to characterize the distributions of stationary period and movement distance. A method to estimate the parameters from empirical data is given, and we apply the model to simulate the movement of two organisms living in different habitats: a stream fish (Cyprinidae: Nocomis leptocephalus) in water, and a root-feeding weevil, Sitona lepidus in the soil. Comparison of the simulations with the measured data shows close agreement. This has an important implication in ecology that the leptokurtic distribution observed at population level does not necessarily mean population heterogeneity as most existing models suggested, in which the population consists of different phenotypes; instead, a homogeneous population moving in homogeneous habitat can also lead to leptokurtic distribution.     

Reducing over-reporting of deterministic co-occurrence patterns in biotic communities

Null models of species co-occurrence are widely used to infer the existence of various ecological processes. Here we investigate the susceptibility of the most commonly used of these models (the C-score in conjunction with the sequential swap algorithm) to type 1 and type 2 errors. To do this we use simulated datasets with a range of numbers of sites, species and coefficients of variation (CV) in species abundance. We find that this model is particularly susceptible to type 1 errors when applied to large matrices and those with low CV in species abundance. As expected, type 2 error rates decrease with increasing numbers of sites and species, although they increase with increasing CV in species abundance. Despite this, power remains acceptable over a wide range of parameter combinations. The susceptibility of this analytical method to type 1 errors indicates that many previous studies may have incorrectly reported the existence of deterministic patterns of species co-occurrence. We demonstrate that in order to overcome the problem of high type 1 error rates, the number of swaps used to generate null distributions for smaller matrices needs to be increased to over 50,000 swaps (well beyond the 5000 commonly used in published analyses and the 30,000 suggested by Lehsten and Harmand, 2006). We also show that this approach reduces type 1 error rates in real datasets. However, even using this solution, larger datasets still suffer from high type 1 error rates. Such datasets therefore require the use of very large numbers of swaps, which calls for improvements in the most commonly used software. In general, users of this powerful analytical method must be aware that they need surprisingly large numbers of swaps to obtain unbiased estimates of structuring in biotic communities.     

Bayesian analysis of abundance for binomial sighting data with unknown number of marked individuals

The mark-resight method for estimating the size of a closed population can in many circumstances be a less expensive and less invasive alternative to traditional mark-recapture. Despite its potential advantages, one major drawback of traditional mark-resight methodology is that the number of marked individuals in the population available for resighting needs to be known exactly. In real field studies, this can be quite difficult to accomplish. Here we develop a Bayesian model for estimating abundance when sighting data are acquired from distinct sampling occasions without replacement, but the exact number of marked individuals is unknown. By first augmenting the data with some fixed number of individuals comprising a marked “super population,” the problem may then be reformulated in terms of estimating the proportion of this marked super population that was actually available for resighting. This then allows the data for the marked population available for resighting to be modeled as random realizations from a binomial logit-normal distribution. We demonstrate the use of our model to estimate the New Zealand robin (Petroica australis) population size in a region of Fiordland National Park, New Zealand. We then evaluate the performance of the proposed model relative to other estimators via a series of simulation experiments. We generally found our model to have advantages over other models when sample sizes are smaller with individually heterogeneous resighting probabilities. Due to limited budgets and the inherent variability between individuals, this is a common occurrence in mark-resight population studies. WinBUGS and R code to carry out these analyses is available from .     

Testing a stochastic version of the Beddington–DeAngelis functional response in foraging shore crabs

Current behaviour-based interference models assume that the predator population is infinitely large and that interference is weak. While the realism of the first assumption is questionable, the second assumption conflicts with the purpose of interference models. Here, we tested a recently developed stochastic version of the Beddington–DeAngelis functional response—which applies to a finite predator population without assuming weak interference—against experimental data of shore crabs (Carcinus maenas) foraging on mussels (Mytilus edulis). We present an approximate maximum likelihood procedure for parameter estimation when only one focal individual is observed, and introduce ‘correction factors’ that capture the average behaviour of the competing but unobserved individuals. We used the method to estimate shore crab handling time, interaction time, and searching rates for prey and competitor. Especially the searching rates were sensitive to variation in prey and competitor density. Incorporating constant parameter values in the model and comparing observed and predicted feeding rates revealed that the predictive power of the model is high. Our stochastic version of the Beddington–DeAngelis model better reflects reality than current interference models and is also amenable for modelling effects of interference on predator distributions.     

Modeling range dynamics in heterogeneous landscapes: invasion of the hemlock woolly adelgid in eastern North America

Range expansion by native and exotic species will continue to be a major component of global change. Anticipating the potential effects of changes in species distributions requires models capable of forecasting population spread across realistic, heterogeneous landscapes and subject to spatiotemporal variability in habitat suitability. Several decades of theory and model development, as well as increased computing power and availability of fine-resolution GIS data, now make such models possible. Still unanswered, however, is the question of how well this new generation of dynamic models will anticipate range expansion. Here we develop a spatially explicit stochastic model that combines dynamic dispersal and population processes with fine-resolution maps characterizing spatiotemporal heterogeneity in climate and habitat to model range expansion of the hemlock woolly adelgid (HWA; Adelges tsugae). We parameterize this model using multiyear data sets describing population and dispersal dynamics of HWA and apply it to eastern North America over a 57-year period (1951-2008). To evaluate the model, the observed pattern of spread of HWA during this same period was compared to model predictions. Our model predicts considerable heterogeneity in the risk of HWA invasion across space and through time, and it suggests that spatiotemporal variation in winter temperature, rather than hemlock abundance, exerts a primary control on the spread of HWA. Although the simulations generally matched the observed current extent of the invasion of HWA and patterns of anisotropic spread, it did not correctly predict when HWA was observed to arrive in different geographic regions. We attribute differences between the modeled and observed dynamics to an inability to capture the timing and direction of long-distance dispersal events that substantially affected the ensuing pattern of spread.     

The good and the bad stopover: behaviours of migrant reed warblers at two contrasting sites

Successful migration for passerine birds depends largely on the quality of stopover habitats, but we still lack complete knowledge of how migrants search for habitats en route and how they behave when landing at poor quality stopover sites. We compared the distance of exploratory movements and stopover durations of the reed warbler Acrocephalus scirpaceus, a reedbed habitat specialist, released at suitable (reed bed) and unsuitable (sand dune) stopover sites. Birds tape-lured during nocturnal migration to a sand dune were captured, radio-tagged, released and tracked at two sites of contrasting habitat quality. Lean birds were found to move further in the dunes (max. 300 m) than in reeds (max. 200 m), whereas ‘fat’ individuals at both sites remained stationary. Birds spent just 1 day in the dunes and up to 13 days in the reeds. Our results suggest that some nocturnal migrants with restricted diurnal exploratory movements depend on stopover site selection when ceasing nocturnal flight.     

Evolving social influence in large populations

Darwinian studies of collective human behaviour, which deal fluently with change and are grounded in the details of social influence among individuals, have much to offer “social” models from the physical sciences which have elegant statistical regularities. Although Darwinian evolution is often associated with selection and adaptation, “neutral” models of drift are equally relevant. Building on established neutral models, we present a general, yet highly parsimonious, stochastic model, which generates an entire family of real-world, right-skew socio-economic distributions, including exponential, winner-take-all, power law tails of varying exponents, and power laws across the whole data. The widely used Barabási and Albert (1999) Science 286: 509-512 “B-A” model of preferential attachment is a special case of this general model. In addition, the model produces the continuous turnover observed empirically within these distributions. Previous preferential attachment models have generated specific distributions with turnover using arbitrary add-on rules, but turnover is an inherent feature of our model. The model also replicates an intriguing new relationship, observed across a range of empirical studies, between the power law exponent and the proportion of data represented in the distribution.     

Feeding and non-feeding aggression can be induced in invasive shore crabs by altering food distribution

Human activity can have a large impact on surrounding ecosystems. For example, humans alter resource distributions for other species, potentially modifying these species competitive dynamics. These changes in local competitive processes are frequently associated with species invasions. Here, we investigate how differences in resource distribution affect competitive behaviour using the highly invasive European shore crab (Carcinus maenas). Using a controlled laboratory experiment in combination with behaviour assays and social network analysis, we show that individuals feeding in habitats with clumped food distributions are more aggressive than individuals feeding in habitats where food is evenly dispersed, and this aggression is present even on days where crabs are not feeding. Additionally, this persistent aggression can be induced, suggesting that individuals of this invasive species possess the flexibility to modify their competitive behaviours in response to differences in food distributions. Furthermore, we show how these individual responses can lead to changes in overall organisation of aggressive interactions within a population. We discuss these results in relation to how human impacts can have long-term effects on competitive behavioural strategies, and how behavioural flexibility can allow invasive species to colonise and persist in highly impacted sites such as urban ecosystems.     

Denning behaviour of the European badger (Meles meles) correlates with bovine tuberculosis infection status

Heterogeneities in behaviours of individuals may underpin important processes in evolutionary biology and ecology, including the spread of disease. Modelling approaches can sometimes fail to predict disease spread, which may partly be due to the number of unknown sources of variation in host behaviour. The European badger is a wildlife reservoir for bovine tuberculosis (bTB) in Britain and Ireland, and individual behaviour has been demonstrated to be an important factor in the spread of bTB among badgers and to cattle. Radio-telemetry devices were deployed on 40 badgers from eight groups to investigate patterns of den (sett) use in a high-density population, where each group had one or two main and three to eight outlier setts in their territory. Badgers were located at their setts for 28 days per season for 1 year to investigate how patterns differed between individuals. Denning behaviour may have a strong influence on contact patterns and the transmission of disease. We found significant heterogeneity, influenced by season, sex and age. Also, when controlling for these, bTB infection status interacting with season was highly correlated with sett use. Test-positive badgers spent more time away from their main sett than those that tested negative. We speculate that wider-ranging behaviour of test-positive animals may result in them contacting sources of infection more frequently and/or that their behaviour may be influenced by their disease status. Measures to control infectious diseases might be improved by targeting functional groups, specific areas or times of year that may contribute disproportionately to disease spread.