Exploring the implications of recreational disturbance on an endangered butterfly using a novel modelling approach

Site and wildlife managers globally are under increasing pressure to implement management strategies that address the negative implications of outdoor recreational activities on wildlife. For many rare and isolated species any anthropogenic activities that cause disturbance could potentially be detrimental to existing populations. Understanding how non-consumptive recreation can influence a species may therefore be critical to its preservation. We developed a novel approach to specifically address this need. Using a combination of field surveys and simulation modelling exercises, we (1) explored the responses of endangered Karner blue butterflies (Lycaeides melissa samuelis) to recreation, (2) assessed whether such responses influenced oviposition rate and/or host plant choice and (3) tested alternative management strategies that could alleviate the negative impacts of recreation. Our field surveys confirmed that Karner blues were sensitive to recreational disturbance. Butterflies flushed at similar speeds and distances from recreationists (2.2 m at 0.17 m/s), as they would from natural threats, such as predators (2.2 m at 0.19 m/s). Incorporating female response parameters into a simulation model revealed that regular disturbance could reduce egg laying potential and significantly restrict host plant choice, which in turn, could impact the butterfly’s population dynamics. However, we established that it was possible to effectively offset the implications of recreational disturbance using our simulation modelling approach. For example, extending Karner blue breeding habitat from trails and other public rights of way has the potential to alleviate such disturbance. Our study demonstrates that the potential impact of recreation on species of conservation concern should not be overlooked.     

Trade-off between human–wildlife conflict risk and recreation conditions

Our study reveals trade-offs associated with human–wildlife conflicts in recreation areas. It is common for recreational users to face risks of wildlife attacks while hiking. Although some studies show that recreational users consider trade-offs between three dimensions of recreational experiences consisting of resource, social and management conditions, there are no studies addressing human–wildlife conflicts from the perspective of the trade-offs for the users. To better understand trade-offs considered by recreational users, we performed a discrete choice experiment. We conducted a questionnaire-based survey among recreational users at the Numameguri hiking trail in Daisetsuzan National Park, Japan, which has a high density of brown bears. Using a series of choice tasks, respondents indicated a preferred trail scenario from a group of alternatives. Scenarios varied in terms of their destinations (resource), crowding (social), management system (management) and bear encounter risk (bear appearances on the trail and bear appearances on mountain slopes seen from the trail). Results show that risk conditions have decisive, negative effects on satisfaction level for those hiking. Specifically, when the risk of encountering brown bears on the trail is high, user satisfaction cannot be increased on the other three dimensions. A consideration of only resource-, social- and management-based indicators is insufficient for understanding recreation experiences in areas with a high risk for human–wildlife conflict. At the recreation areas that are home to large wildlife, a guided tour may leave users more satisfied and safe from wildlife attacks. Understanding trade-offs related to the risk helps tour design and enhances the users’ experience.     

Outdoor recreation causes effective habitat reduction in capercaillie Tetrao urogallus: a major threat for geographically restricted populations

Outdoor recreation inflicts a wide array of impacts on individual animals, many of them reflected in the avoidance of disturbed areas. The scale and spatial extent, however, at which wildlife populations are affected, are mostly unclear. Particularly in geographically isolated populations, where restricted habitat availability may preclude a relocation to undisturbed areas, effective habitat reduction may remain underestimated or even unnoticed, when animals stay in disturbed areas and only show small‐scale responses. Based on telemetry data, we investigated the spatial and seasonal effects of outdoor recreation – in relation to landscape and vegetation conditions – on western capercaillie Tetrao urogallus, considering two scales, home range and within‐home range habitat selection. We determined the distance‐thresholds up to which recreation infrastructures were avoided and estimated the extent of affected habitat for the isolated Black Forest (southwestern Germany) study population. While outdoor recreation did not affect home range selection, strong effects on habitat use within the home range were detected: distance to recreation infrastructure (hiking and cross‐country skiing trails, ski pistes) was the main determinant of habitat selection in winter; in summer, mountain bike trails and hiker's restaurants were avoided up to an average distance of 145 m (CI: 60–1092 m). Around winter‐infrastructure, relative avoidance was recorded up to 320 m (CI: 36–327 m), it was reduced, however, when dense understory provided visual cover. Of the entire population area, between 8–20% (summer) and 8–40% (winter) were affected by outdoor recreation, mainly in the high altitudes. Even without evident large‐scale shifts in species distribution, local‐scale avoidance of outdoor recreation can substantially contribute to effective habitat reduction. Based on our results we recommend a general reduction in recreation infrastructure density in key habitats, the establishment of undisturbed wildlife refuges with a diameter of at least 800 m, as well as enhancing visual protection by maintaining a strip of dense understory along trails.     

Recreationist behaviour in forests and the disturbance of wildlife

Forests are popular locations for outdoor recreation and there is considerable evidence highlighting the positive social impacts of these activities. There is also a body of research outlining the range of potentially negative impacts of recreation on wildlife and habitats. This paper provides a summary of current social and natural scientific knowledge on disturbance caused by walking, cycling, mountain biking, horse riding, off-road vehicles use, camping, and some other recreational activities in forests. We identify more than 40 ecological studies of recreational impacts on forests. Greatest attention has been directed towards walking as an activity and the impacts upon birds, soils and flora although long-term ecological studies of wildlife or habitat disturbance are scarce. Impacts include trampling by foot, hoof and tyre, animal behaviour change and the spread of pests and pathogens. Considerably less work has been carried out on the social dimensions of recreational disturbance. In this article the authors draw on behaviour theory in an attempt to identify the key factors influencing human behaviour in the context of recreational disturbance. Cognitive theories highlight the importance of attitudes and behavioural control, whilst social practice theories emphasise the impact of behavioural routines and contexts. Management actions may be better targeted at promoting alternative behaviours rather than trying to prevent current ‘problem’ behaviours. We advocate greater engagement with these theories to better integrate social science with ecological studies, and improve understanding and management of interactions between recreation needs and conservation.     

Nonmotorized recreation and motorized recreation in shrub‐steppe habitats affects behavior and reproduction of golden eagles (Aquila chrysaetos)

Different forms of outdoor recreation have different spatiotemporal activity patterns that may have interactive or cumulative effects on wildlife through human disturbance, physical habitat change, or both. In western North America, shrub‐steppe habitats near urban areas are popular sites for motorized recreation and nonmotorized recreation and can provide important habitat for protected species, including golden eagles. Our objective was to determine whether recreation use (i.e., number of recreationists) or recreation features (e.g., trails or campsites) predicted golden eagle territory occupancy, egg‐laying, or the probability a breeding attempt resulted in ≥1 offspring (nest survival). We monitored egg‐laying, hatching and fledging success, eagle behavior, and recreation activity within 23 eagle territories near Boise, Idaho, USA. Territories with more off‐road vehicle (ORV) use were less likely to be occupied than territories with less ORV use (β = ?1.6, 85% CI: ?2.8 to ?0.8). At occupied territories, early season pedestrian use (β = ?1.6, 85% CI: ?3.8 to ?0.2) and other nonmotorized use (β = ?3.6, 85% CI: ?10.7 to ?0.3) reduced the probability of egg‐laying. At territories where eagles laid eggs, short, interval‐specific peaks in ORV use were associated with decreased nest survival (β = ?0.5, 85% CI: ?0.8 to ?0.2). Pedestrians, who often arrived near eagle nests via motorized vehicles, were associated with reduced nest attendance (β = ?11.9, 85% CI: ?19.2 to ?4.5), an important predictor of nest survival. Multiple forms of recreation may have cumulative effects on local populations by reducing occupancy at otherwise suitable territories, decreasing breeding attempts, and causing nesting failure. Seasonal no‐stopping zones for motorized vehicles may be an alternative to trail closures for managing disturbance. This study demonstrates the importance of considering human disturbance across different parts of the annual cycle, particularly where multiple forms of recreation have varying spatiotemporal use patterns that create human–wildlife interactions.     

Minimizing animal welfare harms associated with predation management in agro‐ecosystems

The impacts of wild predators on livestock are a common source of human–wildlife conflict globally, and predators are subject to population control for this reason in many situations. Animal welfare is one of many important considerations affecting decisions about predation management. Recent studies discussing animal welfare in this context have presented arguments emphasizing the importance of avoiding intentional harm to predators, but they have not usually considered harms imposed by predators on livestock and other animals. Efforts to mitigate predation impacts (including ‘no control’ approaches) cause a variety of harms to predators, livestock and other wildlife. Successfully minimizing the overall frequency and magnitude of harms requires consideration of the direct, indirect, intentional and unintentional harms imposed on all animals inhabiting agricultural landscapes. We review the harms resulting from the management of dingoes and other wild dogs in the extensive beef cattle grazing systems of Australia to illustrate how these negative impacts can be minimized across both wild and domestic species present on a farm or in a free‐ranging livestock grazing context. Similar to many other predator–livestock conflicts, wild dogs impose intermittent harms on beef cattle (especially calves) including fatal predation, non‐fatal attack (mauling and biting), pathogen transmission, and fear‐ or stress‐related effects. Wild dog control tools and strategies impose harms on dingoes and other wildlife including stress, pain and death as a consequence of both lethal and non‐lethal control approaches. To balance these various sources of harm, we argue that the tactical use of lethal predator control approaches can result in harming the least number of individual animals, given certain conditions. This conclusion conflicts with both traditional (e.g. continuous or ongoing lethal control) and contemporary (e.g. predator‐friendly or no‐control) predation management approaches. The general and transferable issues, approaches and principles we describe have broad applicability to many other human–wildlife conflicts around the world.     

Modeling activity patterns of wildlife using time‐series analysis

The study of wildlife activity patterns is an effective approach to understanding fundamental ecological and evolutionary processes. However, traditional statistical approaches used to conduct quantitative analysis have thus far had limited success in revealing underlying mechanisms driving activity patterns. Here, we combine wavelet analysis, a type of frequency‐based time‐series analysis, with high‐resolution activity data from accelerometers embedded in GPS collars to explore the effects of internal states (e.g., pregnancy) and external factors (e.g., seasonal dynamics of resources and weather) on activity patterns of the endangered giant panda (Ailuropoda melanoleuca). Giant pandas exhibited higher frequency cycles during the winter when resources (e.g., water and forage) were relatively poor, as well as during spring, which includes the giant panda's mating season. During the summer and autumn when resources were abundant, pandas exhibited a regular activity pattern with activity peaks every 24 hr. A pregnant individual showed distinct differences in her activity pattern from other giant pandas for several months following parturition. These results indicate that animals adjust activity cycles to adapt to seasonal variation of the resources and unique physiological periods. Wavelet coherency analysis also verified the synchronization of giant panda activity level with air temperature and solar radiation at the 24‐hr band. Our study also shows that wavelet analysis is an effective tool for analyzing high‐resolution activity pattern data and its relationship to internal and external states, an approach that has the potential to inform wildlife conservation and management across species.     

The direct physical,chemical and biotic impacts on Australian coastal waters due to recreational boating

In economically developed countries it is projected that by around 2015 over 50% of a person’s lifetime will become available for leisure. Demand for leisure needs, already strong, will continue to increase. One segment of the market, outdoor nature-based recreation (including tourism), is growing strongly worldwide. A substantial proportion of these activities are water-based. The associated demand for recreational vessels has increased rapidly in recent years and is projected to continue to trend upwards. Australian trends mirror those internationally. Using Australia as a case study, we review the direct physical, chemical and biotic impacts associated with recreational boating in coastal water environments. Major physical impacts include disturbance due to movement of craft in shallow waters (e.g., turbulence) and the effects of anchoring/drag, noise/interference/collision that impacts on wildlife. The most critical chemical impacts result from pollution due to fuels and oils, defouling treatments (even those not legislated in-country), and human waste (e.g., sewage effluent). Important biotic impacts are the potential continued introduction and secondary spread of non-native species. We conclude that while greater research effort will provide more environmentally benign products, with the increasing popularity of recreation vessels, it will be beyond the resources of Australian governments to police legislation effectively. However, based on Australian’s demonstrated engagement with government in terrestrial environmental management, with their deliberate engagement with the boating fraternity, the impacts of recreational boating would be lessened.     

Behavioral correlates of supplementary feeding of wildlife: Can general conclusions be drawn?

Supplementary feeding is a common, but controversial, tool in wildlife management, because it can benefit both humans and wildlife (e.g., increased wildlife densities), but has certain downsides (e.g., increased disease transmission). For species that are often involved in human-wildlife conflicts, two opposing paradigms with respect to supplementary feeding exist, i.e., (i) that supplementary feeding is efficient to lure animals away from undesired places (i.e., diversionary feeding; hypothesis 1), and (ii) that supplementary feeding stimulates ‘nuisance’ behavior (i.e., increased tolerance for humans and selection for human facilities; hypothesis 2). We formulated an alternative hypothesis (hypothesis 3); i.e., that behavioral variation among individuals dilutes population-wide, general patterns with respect to supplementary feeding. Based on GPS relocation data and resource selection functions, we show that neither of the two opposing management paradigms (hypothesis 1 and 2) hold in a particularly ‘conflict rich’ species, the brown bear (Ursus arctos), because individual variation in selection behavior with respect to supplementary feeding diluted population-wide patterns (hypothesis 3), even under very different environmental contexts (Sweden vs. Slovenia; i.e., different human and bear population density, history and intensity of supplementary feeding, topography, etc.). Our results emphasize that individual variation is an important component of behavioral ecology and should be considered in wildlife management and conservation.     

What's your move? Movement as a link between personality and spatial dynamics in animal populations

Recent studies have established the ecological and evolutionary importance of animal personalities. Individual differences in movement and space‐use, fundamental to many personality traits (e.g. activity, boldness and exploratory behaviour) have been documented across many species and contexts, for instance personality‐dependent dispersal syndromes. Yet, insights from the concurrently developing movement ecology paradigm are rarely considered and recent evidence for other personality‐dependent movements and space‐use lack a general unifying framework. We propose a conceptual framework for personality‐dependent spatial ecology. We link expectations derived from the movement ecology paradigm with behavioural reaction‐norms to offer specific predictions on the interactions between environmental factors, such as resource distribution or landscape structure, and intrinsic behavioural variation. We consider how environmental heterogeneity and individual consistency in movements that carry‐over across spatial scales can lead to personality‐dependent: (1) foraging search performance; (2) habitat preference; (3) home range utilization patterns; (4) social network structure and (5) emergence of assortative population structure with spatial clusters of personalities. We support our conceptual model with spatially explicit simulations of behavioural variation in space‐use, demonstrating the emergence of complex population‐level patterns from differences in simple individual‐level behaviours. Consideration of consistent individual variation in space‐use will facilitate mechanistic understanding of processes that drive social, spatial, ecological and evolutionary dynamics in heterogeneous environments.     

Front Cover

This review highlights the scientific advances concerning the origins of human right‐handedness and language (speech and gestures). The comparative approach we adopted provides evidence that research on human and non‐human animals’ behavioural asymmetries helps understand the processes that lead to the strong human left‐hemisphere specialisation. We review four major non‐mutually exclusive environmental factors that are likely to have shaped the evolution of human and non‐human primates’ manual asymmetry: socioecological lifestyle, postural characteristics, task‐level complexity and tool use. We hypothesise the following scenario for the evolutionary origins of human right‐handedness: the right‐direction of modern humans’ manual laterality would have emerged from our ecological (terrestrial) and social (multilevel system) lifestyle; then, it would have been strengthened by the gradual adoption of the bipedal stance associated with bipedal locomotion, and the increasing level of complexity of our daily tasks including bimanual coordinated actions and tool use. Although hemispheric functional lateralisation has been shaped through evolution, reports indicate that many factors and their mutual intertwinement can modulate human and non‐human primates’ manual laterality throughout their life cycle: genetic and environmental factors, mainly individual sociodemographic characteristics (e.g., age, sex and rank), behavioural characteristics (e.g., gesture per se and gestural sensory modality) and context‐related characteristics (e.g., emotional context and position of target). These environmental (evolutionary and life cycle) factors could also have influenced primates’ manual asymmetry indirectly through epigenetic modifications. All these findings led us to propose the hypothesis of a multicausal origin of human right‐handedness.     

Determinants and co‐expression of anti‐predator responses in amphibian tadpoles: a meta‐analysis

A wide range of taxa respond to perceived predation risk (PPR) through inducible defenses, and many prey are capable of responding both behaviorally and morphologically to the same risk event. In cases where multiple defenses confer protection by independent means (i.e. they are mechanistically independent) responses will either be co‐expressed, or the expression of one defense will limit the capacity (or need) to respond along another axis. Our ability to generate a broad understanding of these patters has been limited, in part, by difficulties in comparing results across studies that employ distinct experimental protocols. Using the extensive literature on tadpole responses to PPR, we conducted a meta‐analysis to identify the ecological and experimental determinants of inducible defence expression. We then assessed whether the magnitude of response to PPR along behavioural versus morphological response axes was positively, or negatively, correlated. The most commonly quantified responses to perceived risk in tadpoles included reductions in movement and swimming behaviour, and altered tail morphology. Our analyses reveal that tadpole behavioural responses are strongly influenced by prey family, predator taxon, evolutionary history with the predator (native versus non‐native), amount of prey consumed by the predator, and how perceived risk was manipulated (e.g. presence versus absence of alarm cues). Tail morphology was similarly influenced by these factors, but also whether the target prey was palatable to predators. Thus, our results identify ecological and experimental features that critically influence the observed effect size in tadpole responses to PPR. A positive correlation between behavioural and morphological responses in studies where both were measured indicates that trait co‐specialization is the predominant pattern of defense deployment in larval amphibians. This positive relationship suggests that survival tends to be maximized in tadpoles through equivalent co‐activation of multiple independent axes of protection, opposed to maximal expression along any single axis. Synthesis Our understanding of plastic responses to perceived predation risk (PPR) has benefited substantially from the vast amount of experimental work examining inducible defences in anuran tadpoles. Indeed this research has illustrated the wide variety of ways that prey animals can respond to the same risk event. We conducted a metaanalysis to identify the key ecological and experimental determinants of inducible defence expression. We then show that, in most cases, behavioural and morphological responses to PPR tend to be co‐expressed suggesting that responding along one axis (moving behaviour) does not limit their ability to respond along another distinct axis (tail morphology).     

Sound the stressor: how Hoatzins (Opisthocomus hoazin) react to ecotourist conversation

Exposure to ecotourists often disrupts animal behavior, which is known to contribute to heightened mortality rates. In the Amazon, the emblematic, communal nesting Hoatzin (Opisthocomus hoazin) is frequently pursued by tourists eager for close views. Such encounters may cause heightened stress levels, and egg or nestling predation due to decreased parental attendance to nests and nestlings. The effect of reducing conversational tourist noise near wildlife is poorly understood, but represents one potential mechanism of mitigating the impacts of ecotourists on wildlife. In this study, we approached Hoatzins by canoe, playing recorded tourist conversations at different volumes. Both the distances from which we observed Hoatzins becoming agitated (e.g., clucking, defecation, etc.) and flush (e.g., flight or climbing away) were positively correlated with volume. Within 10 weeks Hoatzins began to habituate to silent approaches. Tourist conversations, however, continued to elicit the same heightened disturbance responses throughout data collection. Therefore, to have the best chance of seeing Hoatzins at a short distance and minimizing potentially negative disturbances, ecotourists should cease all conversation. Although not tested, silence is probably the best strategy when looking for many wildlife species.     

Fence effects on the behavioural responses of South American fur seals to tourist approaches

Animals that breed in coastal colonies, such as pinnipeds, usually attract tourism, which can negatively affect their resting and breeding behaviour if not managed properly. One strategy to reduce human disturbance is to set up fences, but little is known about their local effectiveness. Our purpose was to assess the behavioural responses of South American fur seals (Arctocephaalus australis) towards tourist approaches before and after the implementation of fences in Cabo Polonio colony (Uruguay). We found that human disturbance levels were similar between years and that the presence of a fence reduced (1) overall fur seal responses to tourists by 60%, (2) the most intense behavioural responses (threat, attack, leaving the colony) by more than half, (3) the responses to large tourist groups (>2 people), which were the most disturbing, (4) the responses to closer (<10 m) tourist approaches, and (5) the responses involving more intrusive tourist behaviours (running, shouting, hand waving). Overall, we showed that after the erection of the fence not only human–wildlife interactions were reduced but also the most stressful fur seal behavioural responses. Although further studies are necessary, our results suggest that the implementation of fences can be a simple and affordable means of minimising human disturbance effects on pinnipeds at local levels (e.g., within colonies), particularly if combined with other strategies (e.g., changes in tourist attitudes).     

How do predators and scavengers locate resource hotspots within a tropical forest?

In many parts of the world, wildlife species congregate at ‘hotspot’ locations that offer feeding opportunities unmatched in the wider landscape. But to exploit those resource‐rich sites, animals must first locate them. In tropical Australia, predators and scavengers (especially dingos, scrub turkeys, snakes, and invasive toads) gather beneath large canopy‐emergent trees that house breeding colonies of metallic starlings (Aplonis metallica). Some wildlife species feed on fallen nestlings whereas others consume the rich insect fauna supported by bird detritus, or the other species attracted to those resources. Those congregations largely cease as soon as colony trees fall, suggesting that wildlife aggregations are responses to bird‐associated cues rather than to specific locations. To identify the proximate cues that elicit congregation of wildlife under such trees, we deployed sound cues (starling‐chatter) and two types of scent cues (soil from beneath a starling tree, and complete nests on broken branches). We recorded visitations by animals with camera‐traps. Starling‐chatter did not attract significant numbers of animals, but soil from beneath colony trees attracted many animals (mostly scrub turkeys). Complete nests attracted nest‐predators (dingos, snakes). Our experiments suggest that faunal aggregations beneath colony trees are driven by proximate responses to distinctive scent cues in the soil, especially for species that obtain their food from that bird‐fertilized substrate; but predators that feed directly on fallen nestlings key in specifically on that resource.     

Behaviour is not Reliably Inferred from End-Product Structure in Caddisflies

Many animals engage in elaborate behavioural sequences in order to build structures. The ‘end‐products’ (e.g., nests of birds, cases of caddisflies) of these behaviours have occasionally been used either to infer details of building behaviour or to infer that structures similar in appearance are constructed by similar behavioural sequences. However, behavioural information can be extrapolated from end‐products only if there is congruence between the end‐product structure and the animal's movements. Case building behaviour was studied in caddisflies (Trichoptera) to examine the relationship between end‐products and behaviour. We found examples of taxa that built similar structures using different behavioural patterns, and taxa that built different structures using similar behaviours, regardless of whether these taxa were closely or distantly related. These findings question the reliability of behavioural inferences based solely on details of their associated end‐products and suggest that end‐product structure should be removed from the definition of animal behaviour.     

Applying an animal‐centric approach to improve ecological restoration

Traditionally, ecological restoration is based on re‐establishing patterns of vegetation communities with the expectation that wildlife will recolonize, restoring the ecological function. However, in many restoration projects, wildlife fails to recolonize, even when vegetation is restored, in many cases because revegetated habitats lack the critical features required by wildlife. We present a new approach to restoration, based on a detailed understanding of ecological process, the mechanisms by which wildlife respond to landscape patterns. Our animal‐centric approach involves measuring the risk‐sensitive decision‐making of individual animals as they balance searching for food, mates, and breeding sites with avoiding being eaten by predators and relates this to fine‐scale habitat and landscape structure. The outcome of these decisions can be measured in occupancy of habitat, the information on which conventional restoration is based. Incorporating landscape genetics allows retrospective assessment of the outcome of dispersal decisions by individual animals on a deeper time frame and at regional scales. Fine‐scale connectivity models can be parameterized with these multiscale spatial and temporal data to direct restoration efforts. We are translating this novel approach to practice in the large Midlands restoration project (4 years, AUD $6 million) in Tasmania, Australia, in partnership with Greening Australia. More than 200 years of intensive agricultural practice in this National Biodiversity Hotspot has resulted in extensive landscape modification, high densities of feral cats, and decline of many native mammals. Our research–practice partnership will alter the way that restoration is done, leading hopefully to successful restoration of wildlife, gene flow, and ecological function.     

Sex, Rank and Age Differences in the Japanese Macaque (Macaca fuscata yakui) Participation in Inter-Group Encounters

In many species interactions among group are often characterized by agonistic behaviour. Although animals may participate in inter‐group encounters in different ways, depending on their energetic requirements, reproductive tactics, and/or developmental stage, the proximate causes affecting an animal's participation in inter‐group encounters are still poorly understood. Indeed, many studies have analysed the behaviour of males and females during inter‐group encounters without considering the importance of additional factors (e.g. rank). This study focuses on wild non‐provisioned Japanese macaques (Macaca fuscata yakui) living on Yakushima Island, Japan. It aims to determine how monkeys of different sex, age, and rank behave during inter‐group encounters and it discusses the implications and consequences of their behaviour on group composition and male dispersal. Males participated significantly more than females in inter‐group encounters, by displaying more aggressive or affiliative behaviour. High‐ranking and/or adult males were more aggressive than low‐ranking and/or subadult males during encounters occurring in the mating season and they also showed more herding behaviour. This trend was not found in inter‐group encounters occurring during the non‐mating season. Finally, males which then emigrated to new groups were low‐ranking and/or subadult individuals. Those males displayed more affiliative behaviour towards foreign males than males which did emigrate. These data indicate that in non‐territorial species with male dominance over female and high competition for mating partners males play an active, and often aggressive, role during inter‐group encounter while female participation is scarce. Factors such as age, rank and period of the year (in seasonally breeding species) have to be taken into considerations when analysing interactions between groups and their effects on group composition and social behaviour.     

A meta‐analysis of human disturbance impacts on Antarctic wildlife

Evidence‐based assessments are increasingly recognized as the best‐practice approach to determine appropriate conservation interventions, but such assessments of the impact of human disturbance on wildlife are rare. Human disturbance comprises anthropogenic activities that are typically non‐lethal, but may cause short‐ and/or longer‐term stress and fitness responses in wildlife. Expanding human activity in the Antarctic region is of particular concern because it increases the scope and potential for increased human disturbance to wildlife in a region that is often thought of as relatively untouched by anthropogenic influences. Here, we use a meta‐analytical approach to synthesise research on human disturbance to wildlife over the last three decades in the Antarctic and sub‐Antarctic region. We combine data from 62 studies across 21 species on the behavioural, physiological and population responses of wildlife to pedestrian, vehicle and research disturbances. The overall effect size indicated a small, albeit statistically significant negative effect of disturbance (?0.39; 95% CI: ?0.60 to ?0.18). Negative effects were found for both physiological and population responses, but no evidence was found for a significant impact on wildlife behavioural responses. Negative effects were found across pedestrian, vehicle and research disturbances. Significant and high among‐study heterogeneity was found in both disturbance and response sub‐groups. Among species, it remains unclear to what extent different forms of disturbance translate into negative population responses. Most current guidelines to limit wildlife disturbance impacts in Antarctica recommend that approaches be tailored to animal behavioural cues, but our work demonstrates that behavioural changes do not necessarily reflect more cryptic, and more deleterious impacts, such as changes in physiology. In consequence, we recommend that pedestrian approach guidelines in the Antarctic region be revisited. Due to the high heterogeneity in effects, management guidelines for different sites and species will need to be developed on a case‐by‐case basis, ideally in conjunction with carefully designed experiments. Guidelines to reduce the impact of research activities per se require development to reduce the potential impacts of conducting research. We identify research questions that, if answered, will further improve the evidence base for guidelines to manage human disturbance in Antarctica.     

Effects of Humans on Behaviour of Wildlife Exceed Those of Natural Predators in a Landscape of Fear

Background

Human disturbance can influence wildlife behaviour, which can have implications for wildlife populations. For example, wildlife may be more vigilant near human disturbance, resulting in decreased forage intake and reduced reproductive success. We measured the effects of human activities compared to predator and other environmental factors on the behaviour of elk (Cervus elaphus Linnaeus 1758) in a human-dominated landscape in Alberta, Canada.

Methodology/Principal Findings

We collected year-round behavioural data of elk across a range of human disturbances. We estimated linear mixed models of elk behaviour and found that human factors (land-use type, traffic and distance from roads) and elk herd size accounted for more than 80% of variability in elk vigilance. Elk decreased their feeding time when closer to roads, and road traffic volumes of at least 1 vehicle every 2 hours induced elk to switch into a more vigilant behavioural mode with a subsequent loss in feeding time. Other environmental factors, thought crucial in shaping vigilance behaviour in elk (natural predators, reproductive status of females), were not important. The highest levels of vigilance were recorded on public lands where hunting and motorized recreational activities were cumulative compared to the national park during summer, which had the lowest levels of vigilance.

Conclusions/Significance

In a human-dominated landscape, effects of human disturbance on elk behaviour exceed those of habitat and natural predators. Humans trigger increased vigilance and decreased foraging in elk. However, it is not just the number of people but also the type of human activity that influences elk behaviour (e.g. hiking vs. hunting). Quantifying the actual fitness costs of human disturbance remains a challenge in field studies but should be a primary focus for future researches. Some species are much more likely to be disturbed by humans than by non-human predators: for these species, quantifying human disturbance may be the highest priority for conservation.