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.     

Why time-limited individuals can make populations more vulnerable to disturbance

Individual variation in disturbance vulnerability (i.e. the likelihood that disturbance negatively affects an individual's fitness) can affect how disturbance impacts animal populations, as even at low disturbance levels some individuals could be severely affected and die. Individual variation in vulnerability can arise due to different responses to disturbance. We propose a new hypothesis that even when individuals respond similarly to disturbance, time-limited individuals are more at risk that their condition deteriorates since they have limited ability to extend their foraging time to compensate for disturbance. We investigate this ‘time-limitation hypothesis' both empirically and mathematically by studying how individuals that differ in their average foraging time (presumably due to differences in foraging efficiency) are affected by disturbance. We used tracking data of 22 wintering Eurasian oystercatchers Haematopus ostralegus to compare time budgets between disturbed and undisturbed tidal periods. In three tidal periods with severe disturbance by transport airplanes, more than a third of the variation in additional flight time and foraging time loss was explained by individual differences. Inefficient individuals that foraged longer in undisturbed tidal periods experienced higher costs in disturbed tidal periods, since they lost more foraging time. We next used an analytical energy balance model to study how time-limited individuals differed in their maximum disturbance thresholds. Both our theoretical model and empirical study suggest that inefficient individuals in a time-limited environment may be unable to increase their foraging time to compensate for the effects of disturbance. Consequently, the magnitude of individual variation in time budgets strongly determines what proportion of the population is at risk that their condition deteriorates due to disturbance. Our hypothesis implies that, when assessing disturbance effects on a population, it is not only important to consider individual variation in disturbance responses, but also variation in time budgets that determine the consequences of disturbance.     

A landscape of coexistence for a large predator in a human dominated landscape

Human related mortality is a major threat for large carnivores all over the world and there is increasing evidence that large predators respond to human related risks in a similar way as prey respond to predation risk. This insight recently led to the conceptual development of a landscape of coexistence that can be used to identify areas which can sustain large predator populations in human dominated landscapes. In this study we applied the landscape of coexistence concept to a large predator in Europe. We investigated to what extent Eurasian lynx Lynx lynx habitat selection is affected by human disturbance in a human dominated landscape. More specifically, we were interested in the existence of a tradeoff between the availability of roe deer, one of their main prey and avoidance of human disturbance and how this affects the spatio‐temporal space use patterns of lynx. We found that lynx face a tradeoff between high prey availability and avoidance of human disturbance and that they respond to this by using areas of high prey availability (but also high human disturbance) during the night when human activity is low. Furthermore our analysis showed that lynx increase their travelling speed and remain more in cover when they are close to areas of high human disturbance. Despite clear behavioral adjustments in response to human presence, prey availability still proved to be the most important predictor of lynx occurrence at small spatial scale, whereas human disturbance was considerably less important. The results of our study demonstrate how spatio‐temporal adaptations in habitat selection enable large carnivores to persist in human dominated landscapes and demonstrate the usefulness of the concept of a landscape of coexistence to develop adaptive management plans for endangered populations of large carnivores.     

A meta-analysis of human disturbance effects on glucocorticoid hormones in free-ranging wild vertebrates

Free-ranging wild vertebrates need to cope with natural and anthropogenic stressors that cause short and/or long-term behavioural and physiological responses. In areas of high human disturbance, the use of glucocorticoid (GC) hormones as biomarkers to measure stress responses is an increasingly common tool for understanding how animals cope with human disturbance. We conducted a meta-analysis to investigate how human disturbances such as habitat conversion, habitat degradation, and ecotourism influence baseline GC hormones of free-ranging wild vertebrates, and we further test the role of protected areas in reducing the impact of such disturbances on these hormones. A total of 58 studies met the inclusion criteria, providing 152 data points for comparing levels of GC hormones under disturbed and undisturbed conditions. The overall effect size suggests that human disturbance does not cause a consistent increase in levels of GC hormones (Hedges' g = 0.307, 95% CI = −0.062 to 0.677). However, when the data were analysed by disturbance type, living in unprotected areas or in areas with habitat conversion were found to increase GC hormone levels compared to living in protected or undisturbed areas. By contrast, we found no evidence that ecotourism or habitat degradation generates a consistent increase in baseline GC hormone levels. Among taxonomic groups, mammals appeared more sensitive to human disturbance than birds. We advocate the use of GC hormones for inferring major human-caused contributors to the stress levels of free-ranging wild vertebrates – although such information needs to be combined with other measures of stress and interpreted in the context of an organism's life history, behaviour, and history of interactions with human disturbance.     

Responses of ants to disturbance in Australia,with particular reference to functional groups

Abstract Ants are widely used as bioindicators in environmental assessment in Australia, partly because the responses of ant communities to disturbance are relatively well understood. In particular, the use of functional groups has provided a predictive framework for analysing ant community responses to disturbance in the absence of reliable information on the responses of individual species. Here we review 45 studies of the responses of Australian ant communities to disturbance, in order to: (i) identify individual species or species‐groups that respond consistently to disturbance; and (ii) examine the usefulness of the functional group scheme as a framework for predicting ant community responses under different disturbance regimes in different biogeographical regions. The most common forms of disturbance in our studies were fire (17 studies), mining (12; mostly studies of minesite restoration) and grazing (7), with other disturbances including clearing, logging, flooding, recreation, urbanization and farming. Responses of individual species were inevitably variable because of differences in vegetation type, severity of disturbance and time since disturbance. However, we identified a range of widespread species that showed predictable responses, including species of the metallica group of Rhytidoponera in temperate Australia (‘increasers’ in relation to disturbance), species of the terebrans and denticulatus groups of Camponotus (increasers), the aeneovirens group of Melophorus (increasers) from the arid zone, and Iridomyrmex pallidus (increaser) from the monsoonal region. The functional group scheme assessed here was not designed specifically in the context of disturbance, but nevertheless in some situations provides a useful framework for analysing ant community responses. Three distinct syndromes of functional group responses can be recognized. First, Dominant Dolichoderinae and Hot Climate Specialists are groups that prefer open environments, and tend to be favoured by low levels of disturbance in well‐forested habitats. Second, Opportunists and often also Generalized Myrmicinae are broadly adapted taxa with wide habitat tolerances, but are particularly sensitive to competitive interactions such that their responses oppose those of Dominant Dolichoderinae. Finally, Cryptic Species and Specialist Predators have highly specialized requirements that make them especially sensitive to disturbance. Functional groups are of most use in situations where disturbance causes substantial change in habitat structure, particularly in the ground‐layer. Functional groups are of least use in very open habitats, where disturbance merely increases what is already extensive bare ground, and has relatively little impact on microclimate. The ant functional group scheme can play an important role in assessing disturbance in mesic Australia, but may be of more limited use for this in the arid zone.     

Text mining for biology - the way forward: opinions from leading scientists

This article collects opinions from leading scientists about how text mining can provide better access to the biological literature, how the scientific community can help with this process, what the next steps are, and what role future BioCreative evaluations can play. The responses identify several broad themes, including the possibility of fusing literature and biological databases through text mining; the need for user interfaces tailored to different classes of users and supporting community-based annotation; the importance of scaling text mining technology and inserting it into larger workflows; and suggestions for additional challenge evaluations, new applications, and additional resources needed to make progress.     

Managing visitor access to seabird colonies: a spatial simulation and empirical observations

Managers of wildlife reserves have a range of tools available to them when considering the best way to provide visitor access while avoiding as many of the negative effects of human disturbance as possible. However, managers lack guidelines as to whether conservation interests are best met by spreading visitors thinly throughout a reserve or by aggregating them in a small area. Here I describe how relationships between disturbance impact and disturbance pressure (the dose–response curve) can be used to address this issue. I generate a spatial simulation of two different models of visitor distribution (one more aggregated than the other) and explicitly model disturbance impact for a variety of dose–response curves. I show that the optimal visitor distribution is likely to depend on the sensitivity of the species and the overall visitor pressure. Importantly, I find that in certain circumstances optimal management can shift from one management option to the other if visitor numbers cross a certain threshold. I use published relationships predicting nesting success of Common Guillemots Uria aalge and Black-legged Kittiwakes Rissa tridactyla to assess optimal management at three nature reserves in Scotland. Optimal management for Guillemots depends on the number of people and the distance between the people and the birds. At sites with high disturbance pressures, management should aim to aggregate visitors in as small an area as possible, whereas at sites with lower disturbance pressure, an even distribution of visitors is favoured. Kittiwake models were not generally accurate, and consequently only site-specific guidelines could be generated, where an even distribution was favoured.     

Seasonal changes in the response of oystercatchers Haematopus ostralegus to human disturbance

The response of foraging animals to human disturbance can be considered as a trade-off between the increased perceived predation risk of tolerating disturbance and the increased starvation risk of not feeding and avoiding disturbance. We show how the response of overwintering oystercatchers Haematopus ostralegus to disturbance is related to their starvation risk of avoiding disturbance. As winter progresses, oystercatcher energy requirements increase and their feeding conditions deteriorate. To survive they spend longer feeding and so have less spare time in which to compensate for disturbance. Later in winter, birds approach a disturbance source more closely and return more quickly after a disturbance. Their behavioural response to disturbance is less when they are having more difficulty surviving and hence their starvation risk of avoiding disturbance is greater. These results have implications for studies which assume that a larger behavioural response means that a species is more vulnerable to disturbance. The opposite may be true. To more fully understand the impact of disturbance, studies should measure both behavioural responses and the ease with which animals are meeting their requirements. Conservation effort should be directed towards species which need to spend a high proportion of their time feeding, but still have a large response to disturbance.     

Human disturbance alters endocrine and immune responses in the Galapagos marine iguana (Amblyrhynchus cristatus)

Anthropogenic disturbance is a relevant and widespread facilitator of environmental change and there is clear evidence that it impacts natural populations. While population-level responses to major anthropogenic changes have been well studied, individual physiological responses to mild disturbance can be equally critical to the long-term survival of a species, yet they remain largely unexamined. The current study investigated the impact of seemingly low-level anthropogenic disturbance (ecotourism) on stress responsiveness and specific fitness-related immune measures in different breeding stages of the marine iguana (Amblyrhynchus cristatus). Specifically, we found stress-induced elevations in plasma corticosterone among tourist-exposed populations relative to undisturbed populations. We also found changes in multiple immunological responses associated with stress-related effects of human disturbance, including bacterial killing ability, cutaneous wound healing, and hemolytic complement activity, and the responses varied according to reproductive state. By identifying health-related consequences of human disturbance, this study provides critical insight into the conservation of a well-known species that has a very distinct ecology. The study also broadens the foundation of knowledge needed to understand the global significance of various levels of human disturbance.     

Human disturbance alters endocrine and immune responses in the Galapagos marine iguana (Amblyrhynchus cristatus)

Anthropogenic disturbance is a relevant and widespread facilitator of environmental change and there is clear evidence that it impacts natural populations. While population-level responses to major anthropogenic changes have been well studied, individual physiological responses to mild disturbance can be equally critical to the long-term survival of a species, yet they remain largely unexamined. The current study investigated the impact of seemingly low-level anthropogenic disturbance (ecotourism) on stress responsiveness and specific fitness-related immune measures in different breeding stages of the marine iguana (Amblyrhynchus cristatus). Specifically, we found stress-induced elevations in plasma corticosterone among tourist-exposed populations relative to undisturbed populations. We also found changes in multiple immunological responses associated with stress-related effects of human disturbance, including bacterial killing ability, cutaneous wound healing, and hemolytic complement activity, and the responses varied according to reproductive state. By identifying health-related consequences of human disturbance, this study provides critical insight into the conservation of a well-known species that has a very distinct ecology. The study also broadens the foundation of knowledge needed to understand the global significance of various levels of human disturbance.     

Cell groups reveal structure of stimulus space

An important task of the brain is to represent the outside world. It is unclear how the brain may do this, however, as it can only rely on neural responses and has no independent access to external stimuli in order to “decode” what those responses mean. We investigate what can be learned about a space of stimuli using only the action potentials (spikes) of cells with stereotyped—but unknown—receptive fields. Using hippocampal place cells as a model system, we show that one can (1) extract global features of the environment and (2) construct an accurate representation of space, up to an overall scale factor, that can be used to track the animal's position. Unlike previous approaches to reconstructing position from place cell activity, this information is derived without knowing place fields or any other functions relating neural responses to position. We find that simply knowing which groups of cells fire together reveals a surprising amount of structure in the underlying stimulus space; this may enable the brain to construct its own internal representations.     

Variability in the area,energy and time costs of wintering waders responding to disturbance

Birds’ responses to human disturbance are interesting due to their similarities to anti‐predator behaviour, and understanding this behaviour has practical applications for conservation management by informing measures such as buffer zones to protect priority species. To understand better the costs of disturbance and whether it will impact on population size, studies should quantify time‐related responses as well as the more commonly reported flight initiation distance (FID). Using waders wintering on an estuarine area, we experimentally disturbed foraging birds on the Wash Embayment, UK, by walking towards them and recording their responses (FID, alert time, time spent in flight, time taken to resume feeding, and total feeding time lost). We present data for 10 species of conservation concern: Curlew Numenius arquata, Oystercatcher Haematopus ostralegus, Bar‐tailed Godwit Limosa lapponica, Grey Plover Pluvialis squatarola, Redshank Tringa totanus, Knot Calidris canutus, Turnstone Arenaria interpres, Ringed Plover Charadrius hiaticula, Sanderling Calidris alba and Dunlin Calidris alpina. Larger species responded more strongly, response magnitude was greater under milder environmental conditions, and responses varied over both small and large spatial scales. The energetic costs of individual responses, however, were low relative to daily requirements and disturbance events were unlikely to be frequent enough to seriously limit foraging time. We suggest, therefore, that wintering wader populations on the Wash are not currently significantly negatively impacted by human disturbance during the intertidal foraging period. This is also likely to be the case at other estuarine sites with comparable access levels, visitor patterns, invertebrate food availability and environmental conditions.     

Impact of human disturbance,density, and environmental conditions on the survival probabilities of pipistrelle bat (Pipistrellus pipistrellus)

Natural and anthropogenic disturbances can strongly impact population dynamics of species and are often responsible for zoonotic emerging infectious diseases. However, long-term studies on the demographic consequences of human disturbances are unusual. We used 6 years (1995–2000) of mark-recapture data to investigate how climatic conditions, human disturbance and density affect sex- and age-specific apparent survival probabilities of the pipistrelle bat (Pipistrellus pipistrellus, Schreber 1774) in a maternity colony. Our study demonstrated that density played an important role in population dynamics of pipistrelle bat and that its effect differed with respect to age and sex. Notably, human disturbance caused a strong decline of adult female survival, suggesting that perturbations have important consequences in bat-colony dynamics. Juvenile female survival was negatively influenced by density, being considerably lower in high densities. In contrast, juvenile and adult males were apparently not affected as they had constant survival probabilities. Although climatic factors can markedly affect population dynamics of temperate insectivorous bats, in this study, the weather conditions did not influence the survival rates of pipistrelle bats. We provide the first report that demonstrates the density-dependent effect on bat survival. That is especially relevant to better understanding of the bat-population dynamics and to evaluate the consequences of human disturbance and their potential changes in the maternity colony structure.     

Tiger and leopard co-occurrence: intraguild interactions in response to human and livestock disturbance

Intraguild interactions have important implications for carnivore demography and conservation. Differences in how predators respond to different forms of disturbance might alter their interaction patterns. We sought to understand how human and livestock disturbance impact co-occurrence of sympatric large carnivores such as tiger (Panthera tigris) and leopard (P. pardus) and thereby mediate the intraguild interaction pattern to enable coexistence of the species in a human-dominated landscape. We surveyed 361 locations in Chitwan National Park, Nepal, to examine how prey abundance and disturbance factors such as human and livestock presence might influence habitat use by tigers and leopards independently and when co-occurring. Single-species single-season models and two-species single-season models were developed to examine hypotheses on unconditional detection and occupancy and species interaction respectively. Pervasive human use of the park had negative impacts on tiger occupancy while the abundance of prey had a positive influence. Despite significant prey overlap between tigers and leopards, none of the native prey species predicted leopard habitat occupancy. However, habitats used extensively by livestock were also used by leopards. Further, we found strong evidence of intraguild competition. For instance tiger occupancy was higher in prey-rich areas and leopard occupancy was low in the sites where tigers were present. These findings, and a species interaction factor of < 1 clearly indicate that leopards avoid tigers, but their use of areas of disturbance enables them to persist in fringe habitats. We provide empirical evidence of how intraguild interaction may result in habitat segregation between competing carnivores, while also showing that human and livestock use of the landscape create disturbance patterns that facilitate co-occurrence of the predators. Thus, because large carnivores compete, some disturbance may mediate coexistence in small protected areas. Understanding such interactions can help address important conservation challenges associated with maintaining diverse carnivore communities in small or disturbed landscapes.     

An experimental assessment of the potential effects of human disturbance on Black Grouse Tetrao tetrix in the North Pennines,England

In the United Kingdom, Black Grouse Tetrao tetrix have declined in range and abundance during the 20th century. In England at present, birds are largely found only on the margins of managed grouse moors in the north, where more than 80% of birds are confined to the North Pennines Area of Outstanding Natural Beauty. The Black Grouse is one of a group of species potentially at risk from increased human recreational disturbance owing to their threatened status and their use of habitats to which a statutory right of human recreational access has recently been granted. To assess the likely impact of increased disturbance to Black Grouse, 77 were caught and radiotagged between 2002 and 2004, and each was randomly assigned to one of three experimental disturbance categories: no disturbance (low), fortnightly disturbance (moderate) or twice weekly disturbance (high). Birds that were disturbed more regularly flushed at greater distances, especially in spring and winter when birds exposed to high disturbance flushed at 32% greater distances than those in moderate disturbance treatments. There were no differences in fecundity (clutch size, hatching success, breeding success) or survival between disturbance treatments. Winter survival was 78% and summer survival 92%. There was no age‐related difference in survival. The disturbance regimes imposed had no discernible impact upon Black Grouse population dynamics. However, in the absence of appropriate data to indicate likely changes in patterns and levels of human recreation resulting from open access, we cannot be sure whether the levels we applied will be representative in the future. Should actual disturbance levels be higher than those we used in this study, we list visitor management options that may help reduce any conflict that arises.     

Habitat type determines the effects of disturbance on the breeding productivity of the Dartford Warbler Sylvia undata

Numerous studies have examined the causes and impacts of human disturbance on birds, but little is known about how these impacts vary among habitats. This is of applied importance both for predicting bird responses to changes in disturbance and in planning how to reduce disturbance impacts. The Dartford Warbler Sylvia undata , a key heathland breeding species, occupies territories in a range of heathland types. Three territory habitat groups were identified: heather-dominated territories, heather territories with significant areas of European Gorse Ulex europaeus and territories containing Western Gorse U. gallii . Productivity was significantly affected by the timing of breeding in all habitats, but disturbance only appeared to have a significant impact on the productivity of birds in heather territories. Disturbance events in heather territories delayed breeding pairs for up to 6 weeks. This significantly decreased both the number of successful broods raised and the average number of chicks fledged per pair. Nests situated close to territory boundaries in heather territories, with high numbers of disturbance events, were more likely to fail outright. It was determined that an average of between 13 and 16 people passing through a heather territory each hour would delay breeding pairs sufficiently to prevent multiple broods.     

Landscape approaches to studying the effects of disturbance on waterbirds

The internationally important populations of waterbirds that winter in the United Kingdom can face intense pressure from human disturbance as a result of the high urbanization found around many protected coastal or inland wetland sites. Here, I describe and evaluate an approach that has been used to investigate the spatial effects of human disturbance on waterbirds. Rather than directly investigating behavioural responses to individual disturbance events, the presence of features in the landscape associated with disturbance is instead used as a surrogate, with the essential aim being to demonstrate that bird numbers or densities are depressed or their behaviour altered in proximity to areas used by humans. This paper first describes case studies that demonstrate the limitations of the basic inference (i.e. that disturbance influences patterns of waterbird distribution or behaviour) and then how investigations might be strengthened. For conclusions to be sound, it is particularly important that other factors, such as food supply, that might also explain the spatial patterns observed are considered or other corroborative evidence presented. The approach is thus least applicable in the most heterogeneous environments where many factors, perhaps spatially autocorrelated, may explain variation in distribution or behaviour. However, greatest confidence in the validity of conclusions may be gained where studies are able to show (ideally by experimental manipulation) that species' distributions or behaviour vary temporally in line with the levels of human use of the features examined. Although its aim and scope are thus limited, the use of a landscape approach, provided that it takes into account other factors affecting spatial variation in bird abundance or behaviour, can provide a preliminary assessment of species avoidance of key sources of disturbance that may offer a framework for more detailed investigation.     

Methods matter: Different biodiversity survey methodologies identify contrasting biodiversity patterns in a human modified rainforest — A case study with amphibians

Understanding how well tropical forest biodiversity can recover following habitat change is often difficult due to conflicting assessments arising from different studies. One often overlooked potentially confounding factor that may influence assessments of biodiversity response to habitat change, is the possibility that different survey methodologies, targeting the same indicator taxon, may identify different patterns and so lead to different conclusions. Here we investigated whether two different but commonly used survey methodologies used to assess amphibian communities, pitfall trapping and nocturnal transects, indicate the same or different responses of amphibian biodiversity to historic human induced habitat change. We did so in a regenerating rainforest study site located in one of the world’s most biodiverse and important conservation areas: the Manu Biosphere Reserve. We show that the two survey methodologies tested identified contrasting biodiversity patterns in a human modified rainforest. Nocturnal transect surveys indicated biodiversity differences between forest with different human disturbance histories, whereas pitfall trap surveys suggested no differences between forest disturbance types, except for community composition. This pattern was true for species richness, diversity, overall abundance and community evenness and structure. For some fine scale metrics, such as species specific responses and abundances of family groups, both methods detected differences between disturbance types. However, the direction of differences was inconsistent between methods. We highlight that for assessments of rainforest recovery following disturbance, survey methods do matter and that different biodiversity survey methods can identify contrasting patterns in response to different types of historic disturbance. Our results contribute to a growing body of evidence that arboreal species might be more sensitive indicators than terrestrial communities.     

Awareness and use of biodiversity collections by fish biologists

A survey of 280 fish biologists from a diverse pool of disciplines was conducted in order to assess the use made of biodiversity collections and how collections can better collect, curate and share the data they have. From the responses, data for how fish biologists use collections, what data they find the most useful, what factors influence the decisions to use collections, how they access the data and explore why some fish biologists make the decision to not use biodiversity collections is collated and reported. The results of which could be used to formulate sustainability plans for collections administrators and staff who curate fish biodiversity collections, while also highlighting the diversity of data and uses to researchers.     

Antipredator behaviour of African ungulates around human settlements

As human populations grow and come into more frequent contact with wildlife, it is important to understand how anthropogenic disturbance alters wildlife behaviour. Using fine‐scale spatial analyses, we examined how proximity to human settlements affects antipredator responses of ungulates. We studied seven common ungulate species (Kirk's dik‐dik, Thomson's gazelle, impala, common warthog, common wildebeest, common zebra and Masai giraffe) in the Tarangire–Manyara ecosystem in northern Tanzania. In zebra and giraffe, flight responses to humans were significantly more likely when closer to settlements; however, there was a weak relationship between flight responses and distance to settlement in all other species. While there was largely a weak relationship between proximity to human settlements, the distribution of settlements in the landscape appears to affect wildlife behaviour, suggesting that animals perceive and respond to spatial variation in risk exerted by humans.