Moving towards acceleration for estimates of activity-specific metabolic rate in free-living animals: the case of the cormorant

1. Time and energy are key currencies in animal ecology, and judicious management of these is a primary focus for natural selection. At present, however, there are only two main methods for estimation of rate of energy expenditure in the field, heart rate and doubly labelled water, both of which have been used with success; but both also have their limitations. 2. The deployment of data loggers that measure acceleration is emerging as a powerful tool for quantifying the behaviour of free-living animals. Given that animal movement requires the use of energy, the accelerometry technique potentially has application in the quantification of rate of energy expenditure during activity. 3. In the present study, we test the hypothesis that acceleration can serve as a proxy for rate of energy expenditure in free-living animals. We measured rate of energy expenditure as rates of O2 consumption (VO2) and CO2 production (VCO2) in great cormorants (Phalacrocorax carbo) at rest and during pedestrian exercise. VO2 and VCO2 were then related to overall dynamic body acceleration (ODBA) measured with an externally attached three-axis accelerometer. 4. Both VO2 and VCO2 were significantly positively associated with ODBA in great cormorants. This suggests that accelerometric measurements of ODBA can be used to estimate VO2 and VCO2 and, with some additional assumptions regarding metabolic substrate use and the energy equivalence of O2 and CO2, that ODBA can be used to estimate the activity specific rate of energy expenditure of free-living cormorants. 5. To verify that the approach identifies expected trends in from situations with variable power requirements, we measured ODBA in free-living imperial cormorants (Phalacrocorax atriceps) during foraging trips. We compared ODBA during return and outward foraging flights, when birds are expected to be laden and not laden with captured fish, respectively. We also examined changes in ODBA during the descent phase of diving, when power requirements are predicted to decrease with depth due to changes in buoyancy associated with compression of plumage and respiratory air. 6. In free-living imperial cormorants, ODBA, and hence estimated VO2, was higher during the return flight of a foraging bout, and decreased with depth during the descent phase of a dive, supporting the use of accelerometry for the determination of activity-specific rate of energy expenditure.     

Assessing the development and application of the accelerometry technique for estimating energy expenditure

A theoretically valid proxy of energy expenditure is the acceleration of an animal's mass due to the movement of its body parts. Acceleration can be measured by an accelerometer and recorded onto a data logging device. Relevant studies have usually derived a measure of acceleration from the raw data that represents acceleration purely due to movement of the animal. This is termed ‘overall dynamic body acceleration’ (ODBA) and to date has proved a robust derivation of acceleration for use as an energy expenditure proxy. Acceleration data loggers are generally easy to deploy and the measures recorded appear robust to slight variation in location and orientation. This review discusses important issues concerning the accelerometry technique for estimating energy expenditure and ODBA; deriving ODBA, calibrating ODBA, acceleration logger recording frequencies, scenarios where ODBA is less likely to be valid, and the power in recording acceleration and heart rate together. While present evidence suggests that ODBA may not quantify energy expenditure during diving by birds and mammals, several recent studies have assessed changes in mechanical work in such species qualitatively through variation in ODBA during periods of submergence. The use of ODBA in field metabolic studies is likely to continue growing, supported by its relative ease of use and range of applications.     

Application of Overall Dynamic Body Acceleration as a Proxy for Estimating the Energy Expenditure of Grazing Farm Animals: Relationship with Heart Rate

Estimating the energy expenditure of farm animals at pasture is important for efficient animal management. In recent years, an alternative technique for estimating energy expenditure by measuring body acceleration has been widely performed in wildlife and human studies, but the availability of the technique in farm animals has not yet been examined. In the present study, we tested the potential use of an acceleration index, overall dynamic body acceleration (ODBA), as a new proxy for estimating the energy expenditure of grazing farm animals (cattle, goats and sheep) at pasture with the simultaneous evaluation of a conventional proxy, heart rate. Body accelerations in three axes and heart rate for cows (n = 8, two breeds), goats (n = 6) and sheep (n = 5) were recorded, and the effect of ODBA calculated from the body accelerations on heart rate was analyzed. In addition, the effects of the two other activity indices, the number of steps and vectorial dynamic body acceleration (VeDBA), on heart rate were also investigated. The results of the comparison among three activity indices indicated that ODBA was the best predictor for heart rate. Although the relationship between ODBA and heart rate was different between the groups of species and breeds and between individuals (P<0.01), the difference could be explained by different body weights; a common equation could be established by correcting the body weights (M: kg): heart rate (beats/min) = 147.263∙M ^-0.141 + 889.640∙M ^-0.179∙ODBA (g). Combining this equation with the previously reported energy expenditure per heartbeat, we estimated the energy expenditure of the tested animals, and the results indicated that ODBA is a good proxy for estimating the energy expenditure of grazing farm animals across species and breeds. The utility and simplicity of the procedure with acceleration loggers could make the accelerometry technique a worthwhile option in field research and commercial farm use.     

Seasonal differences in energy expenditure,flight characteristics and spatial utilization of Dalmatian Pelicans Pelecanus crispus in Greece

Animals typically adjust their behaviour to their changing environment throughout the annual cycle, modulating key processes such as the timing of breeding and the onset of migration. Such behavioural changes are commonly manifested in the movements and the energetic balance of individuals in relation to their species‐specific physiological characteristics, habitat attributes and the environmental properties of their distribution ranges. We used GPS and acceleration data collected using transmitters on free‐ranging birds to quantify annual movement patterns and estimate energy expenditure of the Dalmatian Pelican Pelecanus crispus, a large, soaring avian species which performs short‐distance migration and spends its entire annual cycle in mid‐latitudes. To assess the representativeness of our results, the transmitter effect was also tested. We found that daily trends in the overall dynamic body acceleration (ODBA; a proxy for energy expenditure) differed among seasons, with the highest values occurring during spring and the lowest during winter. Long inter‐lake flights were very rare in winter, and the number of flights and ODBA during spring was higher than during summer, suggesting greater motivation to move in spring. Although transmitters may have affected the birds, as none of the tagged birds bred, we found seasonal differences in behaviour and activity level. The observed patterns in differences in activity levels, long‐distance flights and flight characteristics between seasons suggest an annual rhythm of energy expenditure. These findings allow a better understanding of bird phenology, specifically regarding adaptations to wintering in a cold climate by reducing movement‐driven energy expenditure. Finally, the identification of periods with high and low energy expenditure may guide future conservation efforts by adjusting conservation plans in accordance with changing needs during the annual cycle.     

Dynamic body acceleration increases by 20% during flight ontogeny of greylag geese Anser anser

Despite our knowledge of the biophysical and behavioural changes during flight ontogeny in juvenile birds, little is known about the changes in the mechanical aspects of energy expenditure during early flight development, particularly in migratory species. Here, we investigate in a unique experimental setup how energy expended during flights changes over time beginning with early ontogeny. We calculate overall dynamic body acceleration (ODBA) as a proxy for energy expenditure in a group of hand raised greylag geese Anser anser trained to fly behind a microlight aircraft. We propose two potential hypotheses; energy expenditure either increases with increasing physiological suitability (the ‘physical development hypothesis’), or decreases as a result of behavioural improvements mitigating flight costs (the ‘behavioural development hypothesis’). There was a significant temporal increase of flight duration and ODBA over time, supporting the ‘physical development hypothesis’. This suggests that early on in flight ontogeny behavioural development leading to flight efficiency plays a weaker role in shaping ODBA changes than the increased physical ability to expend energy in flight. We discuss these findings and the implications of flight development on the life history of migratory species.     

Assessing the validity of the accelerometry technique for estimating the energy expenditure of diving double-crested cormorants Phalacrocorax auritus

Over the past few years, acceleration-data loggers have been used to provide calibrated proxies of energy expenditure: the accelerometry technique. Relationships between rate of oxygen consumption and a derivation of acceleration data termed "overall dynamic body acceleration" (ODBA) have now been generated for a range of species, including birds, mammals, and amphibians. In this study, we examine the utility of the accelerometry technique for estimating the energy expended by double-crested cormorants Phalacrocorax auritus to undertake a dive cycle (i.e., a dive and the subsequent pause at the surface before another dive). The results show that ODBA does not calibrate with energy expenditure in diving cormorants, where energy expenditure is calculated from measures of oxygen uptake during surface periods between dives. The possible explanations include reasons why energy expenditure may not relate to ODBA but also reasons why oxygen uptake between dives may not accurately represent energy expenditure during a dive cycle.     

Use of overall dynamic body acceleration for estimating energy expenditure in cormorants: Does locomotion in different media affect relationships?

The way in which animals use and acquire energy is fundamental to their fitness. Overall dynamic body acceleration (ODBA) has recently been suggested as a new method for the determination of energy expenditure in wild animals. Although the relationship between ODBA and energy expenditure has been calibrated using gas-respirometry, it has only been validated for animals moving in a single medium. In this work we examined whether the relationship between ODBA and energy expenditure varies between activity types and in particular, how locomotion in different media affects the regressions using the Imperial Cormorant Phalacrocorax atriceps as a model species. Regressing mean ODBA values for resting, diving and walking periods on a single graph against mean power values, the mass-specific power (W kg⁻¹) was related to ODBA via; Power = 12.09 + 41.31 ODBA (r² = 0.93). Although values for resting, walking and swimming all fell close to a single linear fit, values for flight deviated substantially from this. The different relationships found between locomotory types are discussed in terms of the muscle groups involved in each kind of behavior.     

Measuring Energetics and Behaviour Using Accelerometry in Cane Toads Bufo marinus

Cane toads Bufo marinus were introduced to Australia as a control agent but now have a rapidly progressing invasion front and damage new habitats they enter. Predictive models that can give expansion rates as functions of energy supply and feeding ground distribution could help to maximise control efficiency but to date no study has measured rates of field energy expenditure in an amphibian. In the present study we used the accelerometry technique to generate behavioural time budgets and, through the derivation of ODBA (overall dynamic body acceleration), to obtain estimates of energetics in free ranging cane toads. This represents the first time that accelerometers have been used to not only quantify the behaviour of animals but also assign to those behaviours rates of energy expenditure. Firstly, laboratory calibrations between ODBA and metabolic rate were obtained and used to generate a common prediction equation for the subject toads (R² = 0.74). Furthermore, acceleration data recorded during different behaviours was studied to ascertain threshold values for objectively defining behaviour categories. Importantly, while subsequent accelerometer field deployments were relatively short they agreed with previous studies on the proportion of time that cane toads locomote yet suggest that the metabolic rate of cane toads in the wild may sometimes be considerably higher than might be assumed based on data for other species.     

Space use patterns of mountain hare (<Emphasis Type="Italic">Lepus timidus</Emphasis>) on the Alps

Populations on the limits of species’ distribution can show different behavioral adaptations to strong ecological pressure than in the central part of the range. We investigated space use patterns of alpine mountain hare (Lepus timidus) at two areas on the southern edge of the species’ range. We monitored 34 hares between 2005 and 2008, estimating home range size, overlap, and site fidelity, and compared our results with space use in Scottish and North-European populations. Home ranges of mountain hares did not differ between two study areas with different habitat types. Subadult animals used larger ranges than adults and both age groups reduced home range size in autumn, a period that might be critical for hares due to changes in diet and/or high energy expenditure during the previous breeding season. Home ranges in these alpine populations were smaller than in Scandinavian populations but within the range of populations in different habitat types in Scotland. Seasonal home ranges overlapped considerably, but differed among the sexes: male–female overlap was higher than same sex (male–male and female–female) spatial overlap. Seasonal shifts of home ranges were small, and site fidelity remained high over the seasons, suggesting that resource distribution remained constant throughout the year and that the knowledge of an intensively frequented area is an important element of habitat quality. We concluded that habitat structure and availability of mates interact in affecting mountain hare space use in alpine habitats.     

The relationship between oxygen consumption and body acceleration in a range of species

The ability to measure the energy expenditure of free-ranging animals is of great importance but the techniques available each have their limitations. Recently, as an alternative to more established techniques, an integrated measure of body acceleration termed overall dynamic body acceleration (ODBA) has been used as a calibrated proxy for rate of oxygen consumption (V(O(2))) and hence metabolic rate. The present study tested the potential of this technique, firstly by expanding the range of species for which the V(O(2))-ODBA relationship has been defined and secondly by undertaking a validation exercise to explore the accuracy of predictions made using ODBA. V(O(2))-ODBA relationships during terrestrial locomotion were established for several bipedal and quadrupedal endotherms and compiled with similar relationships previously determined in other species. A model incorporating all of these species showed that ODBA is an excellent predictor of V(O(2)) but there is variation in the V(O(2))-ODBA relationship between species, and further variation within some species. Including measurements such as body mass and structural size in prediction equations might further improve the predictive power of the 'ODBA technique' and eliminate species-specific differences. In the validation exercise, estimate errors were calculated for the species-specific predictive equations. The use of ODBA to estimate V(O(2)) was valid across all species examined and may show a greater potential for estimating energy expenditure for individual animals than other techniques.     

卧龙自然保护区大熊猫空间利用格局动态变化特征

野生动物在长期的栖息地选择过程中,能判断其生境质量,而趋向于选择既能降低能量消耗,又能获得营养价值和能量净收益较高的有利生境。以往的研究多是从统计学方法或宏观尺度对大熊猫生境进行评价,很少考虑到野生动物自身生物学特性及生境选择过程中的空间利用特征。本研究结合家域模型与景观格局分析技术定量分析大熊猫实际空间利用格局的动态变化特征及其破碎化程度,进而反映不同时期大熊猫生境选择模式及栖息地生境适宜性的动态变化。结果表明：卧龙自然保护区大熊猫在对栖息地的实际利用过程中向更适宜的区域集中,使得高适宜等级区域面积有所增大;而边缘生境区域更容易受到自然灾害和人为因素的干扰,破碎化加剧,需要在保护工作中引起足够的重视。从大熊猫行为模式特征出发,在不同时空尺度上,评估大熊猫对生境选择的空间格局变化特征,丰富了野生动物栖息地适宜性评价的时空尺度选择,为更准确地制定保护区管理政策提供了有效的工具。     

Spatiotemporal drivers of energy expenditure in a coastal marine fish

Animal behavior and energy expenditure often vary significantly across the landscape, and quantifying energy expenditure over space and time provides mechanistic insight into ecological dynamics. Yet, spatiotemporal variability in energy expenditure has rarely been explored in fully aquatic species such as fish. Our objective was to quantify spatially explicit energy expenditure for a tropical marine teleost fish, bonefish (Albula vulpes), to examine how bonefish energetics vary across landscape features and temporal factors. Using a swim tunnel respirometer, we calibrated acoustic accelerometer transmitters implanted in bonefish to estimate their metabolic rates and energy expenditure, and applied this technology in situ using a fine-scale telemetry system on a heterogeneous reef flat in Puerto Rico. Bonefish energy expenditure varied most among habitats, with significant interactions between habitat and temporal factors (i.e., diel period, tide state, season). The energy expenditure was generally highest in shallow water habitats (i.e., seagrass and reef crest). Variation in activity levels was the main driver of these differences in energy expenditure, which in shallow, nearshore habitats is likely related to foraging. Bonefish moderate energy expenditure across seasonal fluctuations in temperature, by selectively using shallow nearshore habitats at moderate water temperatures that correspond with their scope for activity. Quantifying how animals expend energy in association with environmental and ecological factors can provide important insight into behavioral ecology, with implications for bioenergetics models.     

Estimating the energy expenditure of free‐ranging polar bears using tri‐axial accelerometers: A validation with doubly labeled water

Measures of energy expenditure can be used to inform animal conservation and management, but methods for measuring the energy expenditure of free‐ranging animals have a variety of limitations. Advancements in biologging technologies have enabled the use of dynamic body acceleration derived from accelerometers as a proxy for energy expenditure. Although dynamic body acceleration has been shown to strongly correlate with oxygen consumption in captive animals, it has been validated in only a few studies on free‐ranging animals. Here, we use relationships between oxygen consumption and overall dynamic body acceleration in resting and walking polar bears Ursus maritimus and published values for the costs of swimming in polar bears to estimate the total energy expenditure of 6 free‐ranging polar bears that were primarily using the sea ice of the Beaufort Sea. Energetic models based on accelerometry were compared to models of energy expenditure on the same individuals derived from doubly labeled water methods. Accelerometer‐based estimates of energy expenditure on average predicted total energy expenditure to be 30% less than estimates derived from doubly labeled water. Nevertheless, accelerometer‐based measures of energy expenditure strongly correlated (r² = 0.70) with measures derived from doubly labeled water. Our findings highlight the strengths and limitations in dynamic body acceleration as a measure of total energy expenditure while also further supporting its use as a proxy for instantaneous, detailed energy expenditure in free‐ranging animals.     

Permissible Home Range Estimation (PHRE) in Restricted Habitats: A New Algorithm and an Evaluation for Sea Otters

Parametric and nonparametric kernel methods dominate studies of animal home ranges and space use. Most existing methods are unable to incorporate information about the underlying physical environment, leading to poor performance in excluding areas that are not used. Using radio-telemetry data from sea otters, we developed and evaluated a new algorithm for estimating home ranges (hereafter Permissible Home Range Estimation, or “PHRE”) that reflects habitat suitability. We began by transforming sighting locations into relevant landscape features (for sea otters, coastal position and distance from shore). Then, we generated a bivariate kernel probability density function in landscape space and back-transformed this to geographic space in order to define a permissible home range. Compared to two commonly used home range estimation methods, kernel densities and local convex hulls, PHRE better excluded unused areas and required a smaller sample size. Our PHRE method is applicable to species whose ranges are restricted by complex physical boundaries or environmental gradients and will improve understanding of habitat-use requirements and, ultimately, aid in conservation efforts.     

Measures of giant panda habitat selection across multiple spatial scales for species conservation

Examining ecological processes across spatial scales is crucial as animals select and use resources at different scales. We carried out field surveys in September 2005, March–September 2006, and April 2007, and used ecological niche factor analysis to determine habitat preferences for the giant panda (Ailuropoda melanoleuca) across 4 spatial scales: daily movement, core range, home range, and seasonal elevational migration. We found that giant pandas prefer conifer forest and mixed forest at higher than average elevation (2,157 m) of study area in the 4 scale models. However, we also observed significant scale differences in habitat selection. The strength of habitat preference increased with scale for the 2 disturbed forests (sparse forest and fragmented forest), and decreased with scale for 0–30° gentle slope and south- and north-facing aspect. Furthermore, habitat suitability patterns were scale-dependent. These findings highlight the need to determine species–environment associations across multiple scales for habitat management and species conservation. © 2012 The Wildlife Society.     

The effect of diet quality on physiological and life-history traits in the harvestman Pachylus paessleri

The balance between energy acquisition and expenditure is critical to the survival and reproductive success of animals. Here we investigate the long-term effects of diet quality on physiological and life-history flexibility in the harvestman, Pachylus paessleri. We used cow meal as a protein-rich diet and potatoes as a carbohydrate-rich diet in order to reproduce two extreme conditions regarding food quality in harvestmen natural habitat. As proxy variables of the energy expenditure process, we quantified standard metabolic rate (maintenance), changes in body mass (somatic condition), and fecundity (reproduction). We found that animals consuming the protein-rich diet were able to increase both their body condition and fecundity. However, the increment in these two life-history traits was correlated with higher maintenance costs. In contrast, the carbohydrate-rich diet did not provide enough specific nutrients for reproductive events, although it may have allowed animals to survive for a long time. Thus, according to the quality of the diet available in the environment, harvestman females can adopt different life-history strategies correlated with phenotypic adjustments at anatomical and physiological levels. In the Mediterranean region, spatial and temporal changes in food quality are typical, so greater phenotypic flexibility is expected to cope with this kind of environmental variation.     

The need for speed: testing acceleration for estimating animal travel rates in terrestrial dead-reckoning systems

Numerous methods are currently available to track animal movements. However, only one of these, dead-reckoning, has the capacity to provide continuous data for animal movements over fine scales. Dead-reckoning has been applied almost exclusively in the study of marine species, in part due to the difficulty of accurately measuring the speed of terrestrial species. In the present study we evaluate the use of accelerometers and a metric known as overall dynamic body acceleration (ODBA) as a proxy for the measurement of speed for use in dead-reckoning. Data were collated from previous studies, for 10 species locomoting on a treadmill and their ODBA measured by an attached data logger. All species except one showed a highly significant linear relationship between speed and ODBA; however, there was appreciable inter- and intra-specific variance in this relationship. ODBA was then used to estimate speed in a simple trial run of a dead-reckoning track. Estimating distance travelled using speed derived from prior calibration for ODBA resulted in appreciable errors. We describe a method by which these errors can be minimised, by periodic ground-truthing (e.g., by GPS or VHF telemetry) of the dead-reckoned track and adjusting the relationship between speed and ODBA until actual known positions and dead-reckoned positions accord.     

基于核密度估计的动物生境适宜度制图方法

生境适宜度制图能提供动物适宜生境的空间分布信息,对野生动物种群管理、保护地规划等非常重要。生境适宜度制图的关键是构建生境适宜度模型(habitat suitability model, HSM),只基于动物出现位置数据构建HSM的方法在实践中得到了非常广泛的应用。然而现有的只基于动物出现位置数据构建HSM的方法还不能很好地直接表达动物生境适宜度和环境因子之间具有生态学意义的数量关系,因此也就不能很好地体现环境因子对动物生境利用的生态学作用。 本文提出了一种基于核密度估计构建HSM的方法,在地理信息系统技术支持下,通过运用核密度估计从代表性的动物出现位置数据中估计出动物出现对各个环境因子的概率密度函数来直接表达生境适宜度与各个环境因子之间的数量关系,以体现环境因子对动物生境利用的生态学作用,在此基础上对生境适宜度与各个环境因子之间的数量关系进行综合构建了具有明确生态学意义的HSM用于动物生境适宜度制图。以美国Voyageures国家公园的白尾鹿(Odocoileus virginianus)生境适宜度制图为例,基于365个出现位置点位数据并结合积雪深度、地表覆被类型、森林边界长度和坡度等环境因子数据,开展了该方法的案例研究。通过交叉验证计算连续Boyce指数对制图结果进行评价,结果表明：基于核密度估计方法构建的HSM预测能力强,所得出的生境适宜度图经10次交叉验证,连续Boyce指数平均值为0.75,标准差为0.11,达到了较高精度。此外,由于基于核密度估计的方法能以“生境适宜度和环境因子之间具有生态学意义的数量关系”的形式来直接体现环境因子对动物生境利用的生态学作用,就模型的可解释性而言,该方法要优于现有的其他构建HSM的方法。     

Simultaneous biologging of heart rate and acceleration, and their relationships with energy expenditure in free-swimming sockeye salmon (Oncorhynchus nerka)

Monitoring the physiological status and behaviour of free-swimming fishes remains a challenging task, although great promise stems from techniques such as biologging and biotelemetry. Here, implanted data loggers were used to simultaneously measure heart rate (f _H), visceral temperature, and a derivation of acceleration in two groups of wild adult sockeye salmon (Oncorhynchus nerka) held at two different water speeds (slow and fast). Calibration experiments performed with individual fish in a swim tunnel respirometer generated strong relationships between acceleration, f _H, tail beat frequency and energy expenditure over a wide range of swimming velocities. The regression equations were then used to estimate the overall energy expenditure of the groups of fish held at different water speeds. As expected, fish held at faster water speeds exhibited greater f _H and acceleration, and correspondingly a higher estimated energy expenditure than fish held at slower water speeds. These estimates were consistent with gross somatic energy density of fish at death, as determined using proximate analyses of a dorsal tissue sample. Heart rate alone and in combination with acceleration, rather than acceleration alone, provided the most accurate proxies for energy expenditure in these studies. Even so, acceleration provided useful information on the behaviour of fish and may itself prove to be a valuable proxy for energy expenditure under different environmental conditions, using a different derivation of the acceleration data, and/or with further calibration experiments. These results strengthen the possibility that biologging or biotelemetry of f _H and acceleration may be usefully applied to migrating sockeye salmon to monitor physiology and behaviour, and to estimate energy use in the natural environment.     

Selecting Habitat to Survive: The Impact of Road Density on Survival in a Large Carnivore

Habitat selection studies generally assume that animals select habitat and food resources at multiple scales to maximise their fitness. However, animals sometimes prefer habitats of apparently low quality, especially when considering the costs associated with spatially heterogeneous human disturbance. We used spatial variation in human disturbance, and its consequences on lynx survival, a direct fitness component, to test the Hierarchical Habitat Selection hypothesis from a population of Eurasian lynx Lynx lynx in southern Norway. Data from 46 lynx monitored with telemetry indicated that a high proportion of forest strongly reduced the risk of mortality from legal hunting at the home range scale, while increasing road density strongly increased such risk at the finer scale within the home range. We found hierarchical effects of the impact of human disturbance, with a higher road density at a large scale reinforcing its negative impact at a fine scale. Conversely, we demonstrated that lynx shifted their habitat selection to avoid areas with the highest road densities within their home ranges, thus supporting a compensatory mechanism at fine scale enabling lynx to mitigate the impact of large-scale disturbance. Human impact, positively associated with high road accessibility, was thus a stronger driver of lynx space use at a finer scale, with home range characteristics nevertheless constraining habitat selection. Our study demonstrates the truly hierarchical nature of habitat selection, which aims at maximising fitness by selecting against limiting factors at multiple spatial scales, and indicates that scale-specific heterogeneity of the environment is driving individual spatial behaviour, by means of trade-offs across spatial scales.