Nils Bunnefeld

A model-driven approach to quantify migration patterns: individual, regional and yearly differences

1. Animal migration has long intrigued scientists and wildlife managers alike, yet migratory species face increasing challenges because of habitat fragmentation, climate change and over-exploitation. Central to the understanding migratory species is the objective discrimination between migratory and nonmigratory individuals in a given population, quantifying the timing, duration and distance of migration and the ability to predict migratory movements. 2. Here, we propose a uniform statistical framework to (i) separate migration from other movement behaviours, (ii) quantify migration parameters without the need for arbitrary cut-off criteria and (iii) test predictability across individuals, time and space. 3. We first validated our novel approach by simulating data based on established theoretical movement patterns. We then formulated the expected shapes of squared displacement patterns as nonlinear models for a suite of movement behaviours to test the ability of our method to distinguish between migratory movement and other movement types. 4. We then tested our approached empirically using 108 wild Global Positioning System (GPS)-collared moose Alces alces in Scandinavia as a study system because they exhibit a wide range of movement behaviours, including resident, migrating and dispersing individuals, within the same population. Applying our approach showed that 87% and 67% of our Swedish and Norwegian subpopulations, respectively, can be classified as migratory. 5. Using nonlinear mixed effects models for all migratory individuals we showed that the distance, timing and duration of migration differed between the sexes and between years, with additional individual differences accounting for a large part of the variation in the distance of migration but not in the timing or duration. Overall, the model explained most of the variation (92%) and also had high predictive power for the same individuals over time (69%) as well as between study populations (74%). 6. The high predictive ability of the approach suggests that it can help increase our understanding of the drivers of migration and could provide key quantitative information for understanding and managing a broad range of migratory species.

Management strategy evaluation: a powerful tool for conservation?

The poor management of natural resources has led in many cases to the decline and extirpation of populations. Recent advances in fisheries science could revolutionize management of harvested stocks by evaluating management scenarios in a virtual world by including stakeholders and by assessing its robustness to uncertainty. These advances have been synthesized into a framework, management strategy evaluation (MSE), which has hitherto not been used in terrestrial conservation. We review the potential of MSE to transform terrestrial conservation, emphasizing that the behavior of individual harvesters must be included because harvester compliance with management rules has been a major challenge in conservation. Incorporating resource user decision-making required to make MSEs relevant to terrestrial conservation will also advance fisheries science.

From migration to nomadism : movement variability in a northern ungulate across its latitudinal range

Understanding the causes and consequences of animal movements is of fundamental biological interest because any alteration in movement can have direct and indirect effects on ecosystem structure and function. It is also crucial for assisting spatial wildlife management under variable environmental change scenarios. Recent research has highlighted the need of quantifying individual variability in movement behavior and how it is generated by interactions between individual requirements and environmental conditions, to understand the emergence of population-level patterns. Using a multi-annual movement data set of 213 individual moose (Alces alces) across a latitudinal gradient (from 56 degrees to 67 degrees N) that spans over 1100 km of varying environmental conditions, we analyze the differences in individual and population-level movements. We tested the effect of climate, risk, and human presence in the landscape on moose movements. The variation in these factors explained the existence of multiple movements (migration, nomadism, dispersal, sedentary) among individuals and seven populations. Population differences were primarily related to latitudinal variation in snow depth and road density. Individuals showed both fixed and flexible behaviors across years, and were less likely to migrate with age in interaction with snow and roads. For the predominant movement strategy, migration, the distance, timing, and duration at all latitudes varied between years. Males traveled longer distances and began migrating later in spring than females. Our study provides strong quantitative evidence for the dynamics of animal movements in response to changes in environmental conditions along with varying risk from human influence across the landscape. For moose, given its wide distributional range, changes in the distribution and migratory behavior are expected under future warming scenarios.

Express your personality or go along with the group: what determines the behaviour of shoaling perch?

Behavioural syndromes, defined as correlated behaviours in different contexts, have been studied across species and taxa including humans as part of a personality concept. While most studies have focused on solitary individuals, less is known on how shoaling fish compromise between own personality and group behaviour. Risk-taking behaviour in 1-year-old perch (Perca fluviatilis) was observed to compare individual behaviour when in a group and when alone. An experimental design gave the fish the choice between foraging in an open area in the presence of a piscivore and hiding in the vegetation. We quantified the variation accountable by the effect of individuals being in a group, individuals alone and repeated measurements, using hierarchical mixed effects models. Within-group variances were low, but when individuals were later tested alone, individual differences explained most of the variation. Still, the individual best linear unbiased predictors (BLUPs) of time spent in the open area, extracted from the random effects of the mixed effects model, were positively correlated with the corresponding BLUPs when alone. The results indicate that individual behavioural traits are to some degree expressed also within groups. Most fish showed a shyer behaviour when alone, but bolder individuals changed less between treatments than did shyer ones, suggesting a more influential role of bold fish in the group.

A Novel Approach to Assessing the Prevalence and Drivers of Illegal Bushmeat Hunting in the Serengeti

Assessing anthropogenic effects on biological diversity, identifying drivers of human behavior, and motivating behavioral change are at the core of effective conservation. Yet knowledge of people’s behaviors is often limited because the true extent of natural resource exploitation is difficult to ascertain, particularly if it is illegal. To obtain estimates of rule-breaking behavior, a technique has been developed with which to ask sensitive questions. We used this technique, unmatched-count technique (UCT), to provide estimates of bushmeat poaching, to determine motivation and seasonal and spatial distribution of poaching, and to characterize poaching households in the Serengeti. We also assessed the potential for survey biases on the basis of respondent perceptions of understanding, anonymity, and discomfort. Eighteen percent of households admitted to being involved in hunting. Illegal bushmeat hunting was more likely in households with seasonal or full-time employment, lower household size, and longer household residence in the village. The majority of respondents found the UCT questions easy to understand and were comfortable answering them. Our results suggest poaching remains widespread in the Serengeti and current alternative sources of income may not be sufficiently attractive to compete with the opportunities provided by hunting. We demonstrate that the UCT is well suited to investigating noncompliance in conservation because it reduces evasive responses, resulting in more accurate estimates, and is technically simple to apply. We suggest that the UCT could be more widely used, with the trade-off being the increased complexity of data analyses and requirement for large sample sizes.

Factors affecting unintentional harvesting selectivity in a monomorphic species

1. Changes in the abundance of populations have always perplexed ecologists but long-term studies are revealing new insights into population dynamic processes. Long-term data are often derived from harvest records although many wild populations face high harvesting pressures leading to overharvesting and extinction. Additionally, harvest records used to describe population processes such as fluctuations in abundance and reproductive success often assume a random off-take. 2. Selective harvesting based on phenotypic characteristics occurs in many species (e.g. trophy hunting, fisheries) and has important implications for population dynamics, conservation and management. 3. In species with no marked morphological differences between the age and sex classes, such as the red grouse Lagopus lagopus scoticus during the shooting season, hunters cannot consciously select for a specific sex or age class during the shooting process but harvest records could still give a biased reflection of the population structure because of differences in behaviour between age and sex classes. 4. This study compared age and sex ratios in the bag with those in the population before shooting for red grouse at different points in the shooting season and different densities, which has rarely been tested before. 5. More young than old grouse were shot at large bag sizes and vice versa for small bag sizes than would be expected from the population composition before shooting. The susceptibility of old males to shooting compared to females increased with bag size and was high at the first time the area was shot but decreased with the number of times an area was harvested. 6. These findings stress that the assumption made in many studies that harvest records reflect the age and sex ratio of the population and therefore reflect productivity can be misleading. 7. In this paper, as in the literature, it is also shown that number of grouse shot reflects grouse density and therefore that hunting selectivity might influence population dynamics in a cyclic species. 8. The study is not only relevant for red grouse but applies to systems showing interactions between selective harvesting and wider ecological processes, such as age- and sex-related parasitism and territoriality, which may drive population fluctuations.

Understanding scales of movement: animals ride waves and ripples of environmental change

Animal movements are the primary behavioural adaptation to spatiotemporal heterogeneity in resource availability. Depending on their spatiotemporal scale, movements have been categorized into distinct functional groups (e.g. foraging movements, dispersal, migration), and have been studied using different methodologies. We suggest striving towards the development of a coherent framework based on the ultimate function of all movement types, which is to increase individual fitness through an optimal exploitation of resources varying in space and time. We developed a novel approach to simultaneously study movements at different spatiotemporal scales based on the following proposed theory: the length and frequency of animal movements are determined by the interaction between temporal autocorrelation in resource availability and spatial autocorrelation in changes in resource availability. We hypothesized that for each time interval the spatiotemporal scales of moose Alces alces movements correspond to the spatiotemporal scales of variation in the gains derived from resource exploitation when taking into account the costs of movements (represented by their proxies, forage availability NDVI and snow depth respectively). The scales of change in NDVI and snow were quantified using wave theory, and were related to the scale of moose movement using linear mixed models. In support of the proposed theory we found that frequent, smaller scale movements were triggered by fast, small-scale ripples of changes, whereas infrequent, larger scale movements matched slow, large-scale waves of change in resource availability. Similarly, moose inhabiting ranges characterized by larger scale waves of change in the onset of spring migrated longer distances. We showed that the scales of movements are driven by the scales of changes in the net profitability of trophic resources. Our approach can be extended to include drivers of movements other than trophic resources (e.g. population density, density of related individuals, predation risk) and may facilitate the assessment of the impact of environmental changes on community dynamics and conservation.

Can supplementary feeding be used to redistribute moose Alces alces

Abstract Foraging patterns, behaviour and the distribution of animals are affected by the availability and distribution of food in the landscape. Increasing numbers of ungulates may also be in conflict with agriculture, timber, infrastructure and conservation interests. Understanding foraging habits of ungulates and how these are affected by a change in forage availability or composition are, therefore, issues of major importance both from ecological and management perspectives. Supplementary feeding (i.e. artificial supply of food) is being used to improve local habitat, and thereby affecting ungulate movements, habitat choice and migration patterns. We experimentally tested the predictions that supplementary feeding redistributes moose Alces alces during two different migration phases (early, i.e. during the onset of migration and late, i.e. in the wintering areas). We used individually marked moose and pellet group counts to investigate the effect of supplementary feeding both at the individual and population level. We monitored 30 moose with GPS-collars before, during and after the supplementary feeding experiment, corresponding to 8-10% of the moose population in two different valleys in Northern Scandinavia. During early migration, moose ignored supplementary feeding sites even though migration routes were close to the sites. At the end of the migration route, supplementary feeding affected moose movement, distribution and behaviour. In conclusion, we suggest that there is a clear difference in response to supplementary feeding by moose due to the phase of migration. We conclude that supplementary feeding can be used under certain conditions to redistribute moose in relation to browsing, or to traffic, preferably at the end-point of migration.

New directions in management strategy evaluation through cross-fertilization between fisheries science and terrestrial conservation

On 1 and 2 June 2010, an international meeting was held at the University of Paris Sud XI, France, organized within the framework of the EU FP7 consortium project HUNT, to bring together fisheries and conservation scientists to discuss a unified framework for the future of management strategies for harvested species.

Being smart about SMART environmental targets

Focus on the negotiation process, not just the end target Global progress toward meeting the Convention on Biological Diversity (CBD) Aichi targets has recently been found wanting (1). The Aichi targets were intended to be SMART (specific, measurable, ambitious, realistic, and time-bound), partly in response to the perception that failure to meet the preceding global biodiversity targets resulted from their lack of SMART-ness (2). Negotiations are building toward the September 2015 United Nations meeting on Sustainable Development Goals (SDGs), which will influence government and business development priorities for decades. Some argue that scientists must engage with the SDG negotiation process to ensure that the environmental targets (e.g., sustainable food production and water-use efficiency) are not vague, modest, or lacking in detailed quantification (3). We caution against focusing only on ensuring that environmental targets are SMART and call for greater attention on the processes that lead to a target being set and met.