Remote techniques for counting and estimating the number of bird–wind turbine collisions at sea: a review

Since the early 1990s, marine wind farms have become a reality, with at least 13 000 offshore wind turbines currently proposed in European waters. There are public concerns that these man-made structures will have a significant negative impact on the many bird populations migrating and wintering at sea. We assess the degree of usefulness and the limitations of different remote technologies for studying bird behaviour in relation to bird–turbine collisions at offshore wind farms. Radar is one of the more powerful tools available to describe the movement of birds in three-dimensional space. Although radar cannot measure bird–turbine collisions directly, it offers the opportunity to quantify input data for collision models. Thermal Animal Detection System (TADS) is an infra red-based technology developed as a means of gathering highly specific information about actual collision rates, and also for parameterizing predictive collision models. TADS can provide information on avoidance behaviour of birds in close proximity to turbine rotor-blades, flock size and flight altitude. This review also assesses the potential of other (some as yet undeveloped) techniques for collecting information on bird flight and behaviour, both pre- and post-construction of the offshore wind farms. These include the use of ordinary video surveillance equipment, microphone systems, laser range finder, ceilometers and pressure sensors.     

Effects of social group size on information transfer and task allocation

Summary Social animals exchange information during social interaction. The rate of interaction and, hence, the rate of information exchange, typically changes with density and density may be affected by the size of the social group. We investigate models in which each individual may be engaged in one of several tasks. For example, the different tasks could represent alternative foraging locations exploited by an ant colony. An individual's decision about which task to pursue depends both on environmental stimuli and on interactions among individuals. We examine how group size affects the allocation of individuals among the various tasks. Analysis of the models shows the following. (1) Simple interactions among individuals with limited ability to process information can lead to group behaviour that closely approximates the predictions of evolutionary optimality models, (2) Because per capita rates of social interaction may increase with group size, larger groups may be more efficient than smaller ones at tracking a changing environment, (3) Group behaviour is determined both by each individual's interaction with environmental stimuli and by social exchange of information. To keep these processes in balance across a range of group sizes, organisms are predicted to regulate per capita rates of social interaction and (4) Stochastic models show, at least in some cases, that the results described here occur even in small groups of approximately ten individuals.     

Predicting the process of extinction in experimental microcosms and accounting for interspecific interactions in single‐species time series

Predicting population extinction risk is a fundamental application of ecological theory to the practice of conservation biology. Here, we compared the prediction performance of a wide array of stochastic, population dynamics models against direct observations of the extinction process from an extensive experimental data set. By varying a series of biological and statistical assumptions in the proposed models, we were able to identify the assumptions that affected predictions about population extinction. We also show how certain autocorrelation structures can emerge due to interspecific interactions, and that accounting for the stochastic effect of these interactions can improve predictions of the extinction process. We conclude that it is possible to account for the stochastic effects of community interactions on extinction when using single‐species time series.     

Modelled sensitivity of avian collision rate at wind turbines varies with number of hours of flight activity input data

Collision risk modelling of birds at wind turbines typically requires vantage point (VP) data to quantify bird flight activity. The number of VP observation hours required to provide such data, and the associated error in predicted collision rate, have not been formally assessed. Using the Band model and a randomization procedure, we examine the sensitivity of collision rate predictions for the White‐tailed Eagle Haliaeetus albicilla to varying hours of input data on flight activity. Variability in collision rate decreased with increasing number of observation hours. However, at the asymptote in variability (about 62 observation hours) there was still considerable variability in predicted collision rate. VP watches are likely to be inherently variable, and collision rate predictions should assess the potential error associated with such results.     

Predictive power of food web models based on body size decreases with trophic complexity

Food web models parameterised using body size show promise to predict trophic interaction strengths (IS) and abundance dynamics. However, this remains to be rigorously tested in food webs beyond simple trophic modules, where indirect and intraguild interactions could be important and driven by traits other than body size. We systematically varied predator body size, guild composition and richness in microcosm insect webs and compared experimental outcomes with predictions of IS from models with allometrically scaled parameters. Body size was a strong predictor of IS in simple modules (r² = 0.92), but with increasing complexity the predictive power decreased, with model IS being consistently overestimated. We quantify the strength of observed trophic interaction modifications, partition this into density‐mediated vs. behaviour‐mediated indirect effects and show that model shortcomings in predicting IS is related to the size of behaviour‐mediated effects. Our findings encourage development of dynamical food web models explicitly including and exploring indirect mechanisms.     

Correlated pay-offs are key to cooperation

The general belief that cooperation and altruism in social groups result primarily from kin selection has recently been challenged, not least because results from cooperatively breeding insects and vertebrates have shown that groups may be composed mainly of non-relatives. This allows testing predictions of reciprocity theory without the confounding effect of relatedness. Here, we review complementary and alternative evolutionary mechanisms to kin selection theory and provide empirical examples of cooperative behaviour among unrelated individuals in a wide range of taxa. In particular, we focus on the different forms of reciprocity and on their underlying decision rules, asking about evolutionary stability, the conditions selecting for reciprocity and the factors constraining reciprocal cooperation. We find that neither the cognitive requirements of reciprocal cooperation nor the often sequential nature of interactions are insuperable stumbling blocks for the evolution of reciprocity. We argue that simple decision rules such as ‘help anyone if helped by someone’ should get more attention in future research, because empirical studies show that animals apply such rules, and theoretical models find that they can create stable levels of cooperation under a wide range of conditions. Owing to its simplicity, behaviour based on such a heuristic may in fact be ubiquitous. Finally, we argue that the evolution of exchange and trading of service and commodities among social partners needs greater scientific focus.     

Collision risk of Montagu’s Harriers Circus pygargus with wind turbines derived from high-resolution GPS tracking

Flight behaviour characteristics such as flight altitude and avoidance behaviour determine the species-specific collision risk of birds with wind turbines. However, traditional observational methods exhibit limited positional accuracy. High-resolution GPS telemetry represents a promising method to overcome this drawback. In this study, we used three-dimensional GPS tracking data including high-accuracy tracks recorded at 3-s intervals to investigate the collision risk of breeding male Montagu's Harriers Circus pygargus in the Dutch–German border region. Avoidance of wind turbines was quantified by a novel approach comparing observed flights to a null model of random flight behaviour. On average, Montagu's Harriers spent as much as 8.2 h per day in flight. Most flights were at low altitude, with only 7.1% within the average rotor height range (RHR; 45–125 m). Montagu's Harriers showed significant avoidance behaviour, approaching turbines less often than expected, particularly when flying within the RHR (avoidance rate of 93.5%). For the present state, with wind farms situated on the fringes of the regional nesting range, collision risk models based on our new insights on flight behaviour indicated 0.6–2.0 yearly collisions of adult males (as compared with a population size of c. 40 pairs). However, the erection of a new wind farm inside the core breeding area could markedly increase mortality (up to 9.7 yearly collisions). If repowering of the wind farms was carried out using low-reaching modern turbines (RHR 36–150 m), mortality would more than double, whereas it would stay approximately constant if higher turbines (RHR 86–200 m) were used. Our study demonstrates the great potential of high-resolution GPS tracking for collision risk assessments. The resulting information on collision-related flight behaviour allows for performing detailed scenario analyses on wind farm siting and turbine design, in contrast to current environmental assessment practices. With regard to Montagu's Harriers, we conclude that although the deployment of higher wind turbines represents an opportunity to reduce collision risk for this species, precluding wind energy developments in core breeding areas remains the most important mitigation measure.     

Individual‐based ecology of coastal birds

Conservation objectives for non‐breeding coastal birds (shorebirds and wildfowl) are determined from their population size at coastal sites. To advise coastal managers, models must predict quantitatively the effects of environmental change on population size or the demographic rates (mortality and reproduction) that determine it. As habitat association models and depletion models are not able to do this, we developed an approach that has produced such predictions thereby enabling policy makers to make evidence‐based decisions. Our conceptual framework is individual‐based ecology, in which populations are viewed as having properties (e.g. size) that arise from the traits (e.g. behaviour, physiology) and interactions of their constituent individuals. The link between individuals and populations is made through individual‐based models (IBMs) that follow the fitness‐maximising decisions of individuals and predict population‐level consequences (e.g. mortality rate) from the fates of these individuals. Our first IBM was for oystercatchers Haematopus ostralegus and accurately predicted their density‐dependent mortality. Subsequently, IBMs were developed for several shorebird and wildfowl species at several European sites, and were shown to predict accurately overwinter mortality, and the foraging behaviour from which predictions are derived. They have been used to predict the effect on survival in coastal birds of sea level rise, habitat loss, wind farm development, shellfishing and human disturbance. This review emphasises the wider applicability of the approach, and identifies other systems to which it could be applied. We view the IBM approach as a very useful contribution to the general problem of how to advance ecology to the point where we can routinely make meaningful predictions of how populations respond to environmental change.     

Distributed cognition and social brains: reductions in mushroom body investment accompanied the origins of sociality in wasps (Hymenoptera: Vespidae)

The social brain hypothesis assumes the evolution of social behaviour changes animals'' ecological environments, and predicts evolutionary shifts in social structure will be associated with changes in brain investment. Most social brain models to date assume social behaviour imposes additional cognitive challenges to animals, favouring the evolution of increased brain investment. Here, we present a modification of social brain models, which we term the distributed cognition hypothesis. Distributed cognition models assume group members can rely on social communication instead of individual cognition; these models predict reduced brain investment in social species. To test this hypothesis, we compared brain investment among 29 species of wasps (Vespidae family), including solitary species and social species with a wide range of social attributes (i.e. differences in colony size, mode of colony founding and degree of queen/worker caste differentiation). We compared species means of relative size of mushroom body (MB) calyces and the antennal to optic lobe ratio, as measures of brain investment in central processing and peripheral sensory processing, respectively. In support of distributed cognition predictions, and in contrast to patterns seen among vertebrates, MB investment decreased from solitary to social species. Among social species, differences in colony founding, colony size and caste differentiation were not associated with brain investment differences. Peripheral lobe investment did not covary with social structure. These patterns suggest the strongest changes in brain investment—a reduction in central processing brain regions—accompanied the evolutionary origins of eusociality in Vespidae.     

Adult predation risk drives shifts in parental care strategies: a long-term study

1. Grouping provides antipredatory benefits, and therefore aggregation tendencies increase under heightened predation risk. In socially breeding groups, however, conflicts over reproductive shares or safety tend to disintegrate groups. Group formation thereby involves a balance between the antipredatory benefits of aggregation and the destabilizing effect of reproductive conflict. 2. We study the grouping behaviour of a facultatively social precocial sea duck with uniparental female care, the eider (Somateria mollissima Linnaeus). Females tend their young solitarily or in groups of 2-5 females. Here, we focus on the effect predation on adults has on group-formation decisions of brood-caring females. 3. By modifying an existing bidding game over care, we model the effects of predation risk on the width of the window of selfishness, which delimits the reproductive sharing allowing cooperation within brood-rearing coalitions, and generate predictions about the relative frequencies of solitary versus cooperative parental care modes. Furthermore, we model the dilution effect as a function of female group size and predation risk. 4. The window of selfishness widens with increasing predation risk, and the dilution of predation risk increases with both female group size and increasing predation risk, yielding the following predictions: (i) cooperative brood care becomes more prevalent and, conversely, solitary brood care less prevalent under heightened predation risk and (ii) group sizes increase concomitantly. 5. We tested these predictions using 13 years of data on female grouping decisions and annual predation rates, while controlling for the potentially confounding effect of female body condition. 6. Our data supported both predictions, where heightened predation risk of nesting females, but not changes in their condition, increased the relative frequency of cooperative brood care. Increased female nesting mortality also resulted in larger groups of cooperative females. 7. The predation risk of incubating females has long-term implications for later parental care decisions. We discuss the potential consequences of predation-induced shifts in group size on per capita fitness and population-wide productivity.     

Phenotypic assortment mediates the effect of social selection in a wild beetle population

Social interactions often have major fitness consequences, but little is known about how specific interacting phenotypes affect the strength of natural selection. Social influences on the evolutionary process can be assessed using a multilevel selection approach that partitions the effects of social partner phenotypes on fitness (referred to as social or group selection) from those of the traits of a focal individual (nonsocial or individual selection). To quantify the contribution of social selection to total selection affecting a trait, the patterns of phenotypic association among interactants must also be considered. We estimated selection gradients on male body size in a wild population of forked fungus beetles (Bolitotherus cornutus). We detected positive nonsocial selection and negative social selection on body size operating through differences in copulation success, indicating that large males with small social partners had highest fitness. In addition, we found that, in low-density demes, the phenotypes of focal individuals were negatively correlated with those of their social partners. This pattern reversed the negative effect of group selection on body size and led to stronger positive selection for body size. Our results demonstrate multilevel selection in nature and stress the importance of considering social selection whenever conspecific interactions occur nonrandomly.     

DRIFT PROMOTES SPECIATION BY SEXUAL SELECTION

Quantitative genetic models of sexual selection have generally failed to provide a direct connection to speciation and to explore the consequences of finite population size. The connection to speciation has been indirect because the models have treated only the evolution of male and female traits and have stopped short of modeling the evolution of sexual isolation. In this article we extend Lande's (1981) model of sexual selection to quantify predictions about the evolution of sexual isolation and speciation. Our results, based on computer simulations, support and extend Lande's claim that drift along a line of equilibria can rapidly lead to sexual isolation and speciation. Furthermore, we show that rapid speciation can occur by drift in populations of appreciable size ( N_e ≥ 1000). These results are in sharp contrast to the opinion of many researchers and textbook writers who have argued that drift does not play an important role in speciation. We argue that drift may be a powerful amplifier of speciation under a wide variety of modeling assumptions, even when selection acts directly on female mating preferences.     

Analytical methods to determine the effective mesoscopic and macroscopic elastic properties of cortical bone

We compare theoretical predictions of the effective elastic moduli of cortical bone at both the meso- and macroscales. We consider the efficacy of three alternative approaches: the method of asymptotic homogenization, the Mori-Tanaka scheme and the Hashin-Rosen bounds. The methods concur for specific engineering moduli such as the axial Young's modulus but can vary for others. In a past study, the effect of porosity alone on mesoscopic properties of cortical bone was considered, taking the matrix to be isotropic. Here, we consider the additional influence of the transverse isotropy of the matrix. We make the point that micromechanical approaches can be used in two alternative ways to predict either the macroscopic (size of cortical bone sample) or mesoscopic (in between micro- and macroscales) effective moduli, depending upon the choice of representative volume element size. It is widely accepted that the mesoscale behaviour is an important aspect of the mechanical behaviour of bone but models incorporating its effect have started to appear only relatively recently. Before this only macroscopic behaviour was addressed. Comparisons are drawn with experimental data and simulations from the literature for macroscale predictions with particularly good agreement in the case of dry bone. Finally, we show how predictions of the effective mesoscopic elastic moduli can be made which retain dependence on the well-known porosity gradient across the thickness of cortical bone.     

The evolution of the social brain: anthropoid primates contrast with other vertebrates

The social brain hypothesis argues that large brains have arisen over evolutionary time as a response to the social and ecological conflicts inherent in group living. We test predictions arising from the hypothesis using comparative data from birds and four mammalian orders (Carnivora, Artiodactyla, Chiroptera and Primates) and show that, across all non-primate taxa, relative brain size is principally related to pairbonding, but with enduring stable relationships in primates. We argue that this reflects the cognitive demands of the behavioural coordination and synchrony that is necessary to maintain stable pairbonded relationships. However, primates differ from the other taxa in that they also exhibit a strong effect of group size on brain size. We use data from two behavioural indices of social intensity (enduring bonds between group members and time devoted to social activities) to show that primate relationships differ significantly from those of other taxa. We suggest that, among vertebrates in general, pairbonding represents a qualitative shift from loose aggregations of individuals to complex negotiated relationships, and that these bonded relationships have been generalized to all social partners in only a few taxa (such as anthropoid primates).     

Social Selection and Indirect Genetic Effects in Structured populations

Social selection and indirect genetic effects (IGEs) are established concepts in both behavioural ecology and evolutionary genetics. While IGEs describe effects of an individual’s genotype on phenotypes of social partners (and may thus affect their fitness indirectly), the concept of social selection assumes that a given phenotype in one individual affects the fitness of other individuals directly. Although different frameworks, both have been used to investigate the evolution of social traits, such as cooperative behaviour. Despite their similarities (both concepts consider interactions among individuals), they differ in the type of interaction. It remains unclear whether the two concepts make the same predictions about evolutionary trajectories or not. To address this question, we investigate four possible scenarios of social interactions and compare the effects of IGEs and social selection for trait evolution in a multi-trait multi-member model. We show that the two mechanisms can yield similar evolutionary outcomes and that both can create selection pressure at the group level. However, the effect of IGEs can be stronger due to the possibility of feedback loops. Finally, we demonstrate that IGEs, but not social selection gradients, may lead to differences in the direction of evolutionary response between genotypes and phenotypes.     

Cannibalism as an interacting phenotype: precannibalistic aggression is influenced by social partners in the endangered Socorro Isopod (Thermosphaeroma thermophilum)

Models for the evolution of cannibalism highlight the importance of asymmetries between individuals in initiating cannibalistic attacks. Studies may include measures of body size but typically group individuals into size/age classes or compare populations. Such broad comparisons may obscure the details of interactions that ultimately determine how socially contingent characteristics evolve. We propose that understanding cannibalism is facilitated by using an interacting phenotypes perspective that includes the influences of the phenotype of a social partner on the behaviour of a focal individual and focuses on variation in individual pairwise interactions. We investigated how relative body size, a composite trait between a focal individual and its social partner, and the sex of the partners influenced precannibalistic aggression in the endangered Socorro isopod, Thermosphaeroma thermophilum. We also investigated whether differences in mating interest among males and females influenced cannibalism in mixed sex pairs. We studied these questions in three populations that differ markedly in range of body size and opportunities for interactions among individuals. We found that relative body size influences the probability of and latency to attack. We observed differences in the likelihood of and latency to attack based on both an individual's sex and the sex of its partner but found no evidence of sexual conflict. The instigation of precannibalistic aggression in these isopods is therefore a property of both an individual and its social partner. Our results suggest that interacting phenotype models would be improved by incorporating a new conditional ψ, which describes the strength of a social partner's influence on focal behaviour.     

Many eyes or many ewes: vigilance tactics in female bighorn sheep Ovis canadensis vary according to reproductive status

In gregarious animals, there is usually a negative relationship between individual vigilance and group size. This effect of group size is generally explained by increasing probability of predator detection (the many-eyes hypothesis) and by the dilution of risk occurring in larger groups. Few studies have attempted to examine the specific implications of either hypothesis on the expected vigilance pattern of an animal. Here we examine whether reproductive status affects vigilance patterns in bighorn sheep Ovis canadensis ewes. We also test whether the observed vigilance patterns are consistent with predictions from dilution or detection models of vigilance. Although vigilance decreased with increasing group size, vigilance tactics differed between barren and lactating females. Lactating ewes relied solely on predator detection. In contrast, barren ewes benefited from both detection and dilution effects when group size increased and adjusted vigilance effort according to the proportion of lactating ewes in their group. It is generally assumed that gregariousness increases safety. Here we further show that reproductive status influenced how animals reduce predation risk and that some individuals take advantage of the vigilance effort provided by others.     

Nonadditive effects of group membership can lead to additive group phenotypes for anti-predator behaviour of guppies, Poecilia reticulata

Nonadditive effects of group membership are generated when individuals respond differently to the same social environment and may alter predictions about how behavioural evolution will occur. Despite this importance, the relationship between an individual's behaviour in two different social contexts and how reciprocal interactions among individuals within groups influence group behaviour are poorly understood. Guppy anti-predator behaviour can be used to explore how individuals behaviourally respond to changes in social context. Individuals from two strains were tested for response to a model predator alone and in groups to evaluate how individuals alter their behaviour in response to social context and how group phenotype relates to individual behaviour. Nonadditive effects of group membership were detected for a number of behaviours, revealing that the effect of being in a group differed among individuals. These nonadditive effects, however, yielded an additive group phenotype. That is, the average behaviour of the group was equal to the average of its parts, for all behaviours in both strains. Such an additive group phenotype may have resulted because all individuals within a group respond to the specific social environment provided by the other members of their group.     

Collision risk and micro-avoidance rates of birds with wind turbines in Flanders

Capsule Local factors can lead to strong variation in mortality rate and collision risk that obscures possible effects of turbine size in wind farms.

Aims The impact of bird collisions was studied at eight land-based wind farm sites with a total of 66 small to large turbines in order to assess the mortality rate and collision risk.

Methods Searches for collision fatalities were performed under all turbines with a minimum search interval of 14 days. Mortality rate was calculated with corrections for available search area, scavenging and search efficiency. Flight movements of birds crossing five of the wind farm sites were recorded during a minimum of four days per site. Actual collision risk was then calculated as the number of collision fatalities relative to the average surveyed flight intensity.

Results Mortality rate was 21 birds per turbine per year on average. Most fatalities were local common species (e.g. gulls) but rarer species were also found (e.g. terns, raptors and waders). Collision risk of gulls was 0.05% and 0.08% on average for birds, respectively, flying at turbine and rotor height through the wind farms (0.09% and 0.14% maximum). Large gulls had a significant higher collision risk than small gulls at rotor height. Mortality rate and collision risk were not significantly related to turbine size. The results were integrated in a widely used collision risk model to obtain information of micro-avoidance, i.e. the proportion of birds that fly through the wind farm but avoid passing through the rotor swept area of the turbines. For gulls, this micro-avoidance was 96.1% and 96.3% on average for birds, respectively, flying at turbine and rotor height through the wind farms.

Conclusion The results indicate that local factors can lead to strong variation in mortality rate and collision risk that obscures possible effects of turbine size in wind farms. However, large turbines have more installed capacity (MW), so repowering wind farms with larger but fewer wind turbines, could reduce total mortality at certain locations.