BACKGROUND The movement of animals makes them fascinating but difficult study subjects. Animal movements underpin many biological phenomena, and understanding them is critical for applications in conservation, health, and food. Traditional approaches to animal tracking used field biologists wielding antennas to record a few dozen locations per animal, revealing only the most general patterns of animal space use. The advent of satellite tracking automated this process, but initially was limited to larger animals and increased the resolution of trajectories to only a few hundred locations per animal. The last few years have shown exponential improvement in tracking technology, leading to smaller tracking devices that can return millions of movement steps for ever-smaller animals. Finally, we have a tool that returns high-resolution data that reveal the detailed facets of animal movement and its many implications for biodiversity, animal ecology, behavior, and ecosystem function. ADVANCES Improved technology has brought animal tracking into the realm of big data, not only through high-resolution movement trajectories, but also through the addition of other on-animal sensors and the integration of remote sensing data about the environment through which these animals are moving. These new data are opening up a breadth of new scientific questions about ecology, evolution, and physiology and enable the use of animals as sensors of the environment. High–temporal resolution movement data also can document brief but important contacts between animals, creating new opportunities to study social networks, as well as interspecific interactions such as competition and predation. With solar panels keeping batteries charged, “lifetime” tracks can now be collected for some species, while broader approaches are aiming for species-wide sampling across multiple populations. Miniaturized tags also help reduce the impact of the devices on the study subjects, improving animal welfare and scientific results. As in other disciplines, the explosion of data volume and variety has created new challenges and opportunities for information management, integration, and analysis. In an exciting interdisciplinary push, biologists, statisticians, and computer scientists have begun to develop new tools that are already leading to new insights and scientific breakthroughs. OUTLOOK We suggest that a golden age of animal tracking science has begun and that the upcoming years will be a time of unprecedented exciting discoveries. Technology continues to improve our ability to track animals, with the promise of smaller tags collecting more data, less invasively, on a greater variety of animals. The big-data tracking studies that are just now being pioneered will become commonplace. If analytical developments can keep pace, the field will be able to develop real-time predictive models that integrate habitat preferences, movement abilities, sensory capacities, and animal memories into movement forecasts. The unique perspective offered by big-data animal tracking enables a new view of animals as naturally evolved sensors of environment, which we think has the potential to help us monitor the planet in completely new ways. A massive multi-individual monitoring program would allow a quorum sensing of our planet, using a variety of species to tap into the diversity of senses that have evolved across animal groups, providing new insight on our world through the sixth sense of the global animal collective. We expect that the field will soon reach a transformational point where these studies do more than inform us about particular species of animals, but allow the animals to teach us about the world.

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Terrestrial animal tracking as an eye on life and planet

Phenotype-based selective harvests, including trophy hunting, can have important implications for sustainable wildlife management if they target heritable traits. Here we show that in an evolutionary response to sport hunting of bighorn trophy rams (Ovis canadensis) body weight and horn size have declined significantly over time. We used quantitative genetic analyses, based on a partly genetically reconstructed pedigree from a 30-year study of a wild population in which trophy hunting targeted rams with rapidly growing horns, to explore the evolutionary response to hunter selection on ram weight and horn size. Both traits were highly heritable, and trophy-harvested rams were of significantly higher genetic 'breeding value' for weight and horn size than rams that were not harvested. Rams of high breeding value were also shot at an early age, and thus did not achieve high reproductive success. Declines in mean breeding values for weight and horn size therefore occurred in response to unrestricted trophy hunting, resulting in the production of smaller-horned, lighter rams, and fewer trophies.

Undesirable evolutionary consequences of trophy hunting

1. Although the home range is a fundamental ecological concept, there is considerable debate over how it is best measured. There is a substantial literature concerning the precision and accuracy of all commonly used home range estimation methods; however, there has been considerably less work concerning how estimates vary with sampling regime, and how this affects statistical inferences. 2. We propose a new procedure, based on a variance components analysis using generalized mixed effects models to examine how estimates vary with sampling regime. 3. To demonstrate the method we analyse data from one study of 32 individually marked roe deer and another study of 21 individually marked kestrels. We subsampled these data to simulate increasingly less intense sampling regimes, and compared the performance of two kernel density estimation (KDE) methods, of the minimum convex polygon (MCP) and of the bivariate ellipse methods. 4. Variation between individuals and study areas contributed most to the total variance in home range size. Contrary to recent concerns over reliability, both KDE methods were remarkably efficient, robust and unbiased: 10 fixes per month, if collected over a standardized number of days, were sufficient for accurate estimates of home range size. However, the commonly used 95% isopleth should be avoided; we recommend using isopleths between 90 and 50%. 5. Using the same number of fixes does not guarantee unbiased home range estimates: statistical inferences differ with the number of days sampled, even if using KDE methods. 6. The MCP method was highly inefficient and results were subject to considerable and unpredictable biases. The bivariate ellipse was not the most reliable method at low sample sizes. 7. We conclude that effort should be directed at marking more individuals monitored over long periods at the expense of the sampling rate per individual. Statistical results are reliable only if the whole sampling regime is standardized. We derive practical guidelines for field studies and data analysis.

https://besjournals.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1365-2656.2006.01164.x

Effects of sampling regime on the mean and variance of home range size estimates

Summary
1Nutrition influences most aspects of animal ecology: juvenile growth rates and adult mass gain, body condition, probability of pregnancy, over-winter survival, timing of parturition, and neonatal birth mass and survival. We provide an overview among ungulates of the extent of these influences resulting from interactions among bioenergetics, foraging, and nutritional demands.
2 Body condition of an animal is the integrator of nutritional intake and demands, affecting both survival and reproduction. The deposition and mobilization of body fat and body protein vary with physiological requirements and environmental conditions as species use dietary income and body stores to integrate the profits of summer and the demands of winter. Results from our simulation model and uncertainty analysis of the influence of body mass and changes in body composition of Rangifer over winter indicate that percent body fat rather than body mass in early winter is most important in determining whether animals die, live without reproducing, or live and reproduce. Animal responses are also sensitive to rates of change in body protein. Depending on timing of calving and maternal reserves, seasonal habitats vary in their nutritional value for the production of offspring.
3For free-ranging animals, life is a balance among numerous ecological factors, including nutritional requirements, nutritional resources to meet those demands, and intra- and inter-specific interactions. Predation effects on population demography may mask nutritional limitations of habitat. We suggest that over the long term of life histories, ungulates use seasonal strategies that minimize the maximum detriment, and that the basis for most strategies is nutritional.

https://besjournals.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-2435.2009.01528.x

Nutrition integrates environmental responses of ungulates.

It is unclear when, where and how novel pathogens such as human immunodeficiency virus (HIV), monkeypox and severe acute respiratory syndrome (SARS) will cross the barriers that separate their natural reservoirs from human populations and ignite the epidemic spread of novel infectious diseases. New pathogens are believed to emerge from animal reservoirs when ecological changes increase the pathogen's opportunities to enter the human population1 and to generate subsequent human-to-human transmission2. Effective human-to-human transmission requires that the pathogen's basic reproductive number, R0, should exceed one, where R0 is the average number of secondary infections arising from one infected individual in a completely susceptible population3. However, an increase in R0, even when insufficient to generate an epidemic, nonetheless increases the number of subsequently infected individuals. Here we show that, as a consequence of this, the probability of pathogen evolution to R0 > 1 and subsequent disease emergence can increase markedly.

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The role of evolution in the emergence of infectious diseases

It has been suggested that patterns of habitat selection of animals across spatial scales should reflect the factors limiting individual fitness in a hierarchical fashion. Animals should thus select habitats that permit avoidance of the most important limiting factor at large spatial scales while the influence of less important factors should only be evident at fine scales. We tested this hypothesis by investigating moose Alces alces habitat selection using GPS telemetry in an area where the main factors limiting moose numbers were likely (in order of decreasing importance) predation risk, food availability and snow. At the landscape scale, we predicted that moose would prefer areas where the likelihood of encountering wolves was low or areas where habitats providing protection from predation were dominant. At the home-range scale, we predicted that moose selection would be driven by food availability and snow depth. Wolf territories were delineated using telemetry locations and the study area was divided into 3 sectors that differed in terms of annual snowfall. Vegetation surveys yielded 6 habitat categories that differed with respect to food availability, and shelter from predation or snow. Our results broadly supported the hypothesis because moose reacted to several factors at each scale. At the landscape scale, moose were spatially segregated from wolves by avoiding areas receiving the lowest snowfall, but they also preferentially established their home range in areas where shelter from snow bordered habitat types providing abundant food. At the home-range scale, moose also traded off food availability with avoidance of deep snow and predation risk. During winter, moose increased use of stands providing shelter from snow along edges with stands providing abundant food. Habitat selection patterns of females with calves differed from that of solitary moose, the former being associated primarily with habitats providing protection from predation. Animals should attempt to minimize detrimental effects of the main limiting factors when possible at the large scale. However, when the risk associated with several potential limiting factors varies with scale, we should expect animals to make trade-offs among these.

Linking moose habitat selection to limiting factors

Home-range size is widely used in wildlife biology to assess animal-habitat relationships. But home-range size estimation largely depends on the estimator and sample size used. Using 3 different estimators (minimum convex polygons, fixed-kernels, clusters), we used data collected on moose (Alces alces), fitted with Global Positioning System (GPS) collars (which allow automatic recording of large data sets with fewer field constraints than VHF telemetry) to investigate to what extent increasing the number of locations affects home-range size estimations. Our results indicated that 100 to 300 locations per animal annually and 30 to 100 locations seasonally were needed to reach an asymptote. High biases occurred below this asymptotic value. Minimum convex polygons consistently underestimated home-range sizes, but fixed-kernel and cluster estimators followed variable trends that often overestimated home-range sizes. Low sampling efforts also affected the number of animals needed to ensure an adequate statistical power of analysis to compare space use between 2 groups. Despite the higher cost, our findings indicate that in most instances, GPS telemetry is better suited than conventional radiotelemetry to estimate home-range sizes precisely and accurately. Moreover, this tracking technique is not limited by meteorological constraints and allows for the collection of similar sample sizes for all tracked animals, which is of major importance for further comparisons of space use among individuals.

http://www.jstor.org/stable/pdfplus/3802962.pdf

Effects of sampling effort based on GPS telemetry on home-range size estimations

Forest harvesting involves the creation of roads and cutblocks, both of which can influence animal habitat use. We evaluated the cumulative effects of forestry on habitat selection by six packs of gray wolf (Canis lupus) widely distributed in Quebec’s boreal forest. Resource selection functions were used to evaluate cumulative effects at two levels. First, we studied how the response of wolves to roads and cutblocks varied within their home range (HR level) as a function of the local abundance of these habitat features. Second, we assessed whether differences in the response to roads and cutblocks observed among packs (inter-HR level) could be explained by variations in their average abundance among individual home ranges. At the HR level, we found that cumulative effects shaped habitat selection of wolves, and the nature of the effects varied during the year. For example, we detected a decrease in the selection of roads following an increase in local road density during the rendez-vous and the nomadic periods, but not during the denning period. At the inter-HR level, we found a functional response to logging activity only during the denning period. Packs with home ranges characterized by a larger proportion of recent cutblocks selected these cutblocks more strongly. We conclude that cumulative effects of logging activities occur at multiple levels, and these effects can have profound effects on habitat use by wolves, thereby influencing spatial predator–prey dynamics. Wildlife conservation and management in boreal ecosystems should thus account for cumulative impacts of anthropogenic features on animal distribution.

Cumulative effects of forestry on habitat use by gray wolf (Canis lupus) in the boreal forest

:Among ungulate species living in boreal regions, moose (Alces alces) are most likely to suffer from heat stress since they have a relatively low upper critical temperature (14 °C in summer and -5 °C in winter). We tested the hypothesis that moose adopt behaviours to reduce thermoregulatory costs. We predicted that moose exposed to high intensities of solar radiation or high air temperatures would 1) seek a type of vegetation association that provides thermal shelter and 2) reduce activity. We also predicted that these behaviours would be most evident in summer. Thermal-shelter use and activity of 30 free-ranging moose were measured over 3 y in a 940-km2 study area of boreal forest using GPS telemetry collars. The effect of solar radiation and air temperature on thermal-shelter use and activity were assessed using logistic and multiple regression analyses. Habitat use and activity rates of moose were related to air temperature but not solar radiation. The probability of finding moose in thermal sh...

Behavioural responses of moose to thermal conditions in the boreal forest

Roads often negatively affect terrestrial wildlife, via habitat loss or fragmentation, noise, and direct mortality. We studied moose (Alces alces) behavior relative to a road network, in an area with a history of moose-vehicle accidents, to determine when moose were crossing roadways or using areas near roads and to investigate if environmental factors were involved in this behavior. We tracked 47 adult moose with Global Positioning System collars in a study area crossed by highways and forest roads. We hypothesized that moose would avoid crossing roads but would make occasional visits to roadsides to feed on sodium-rich vegetation and avoid biting insects. Further, we expected moose avoidance to be greater for highways than forest roads. We recorded 196,710 movement segments but only observed 328 highway and 1,172 forest-road crossings (16 and 10 times lower than expected by chance). Moose usually avoided road proximity up to ≥500 m on each side but 20% of collared moose made visits to areas within 50 m of highways, which might have resulted from moose searching for sodium in vegetation and roadside salt pools. In fact, vegetation along highways had higher sodium concentrations and was browsed in similar proportions to vegetation in adjacent forest, despite moose avoidance of these zones. Moose, however, did not use areas near roads more during periods of biting insect abundance. Our results supported the hypothesis of scale-dependent selection by moose; avoidance of highways at a coarse scale may confer long-term benefits, whereas selection of highway corridors at finer scales may be part of a strategy to overcome short-term limiting factors such as sodium deficiency. We found a positive relationship between home-range size and the proportion of road axes they contained, suggesting that moose either compensated for habitat loss or made specific movements along highways to gather sodium. The presence of sodium along highways likely increases moose-vehicle accident risks. Removal of salt pools or use of a de-icing salt other than sodium chloride should render highway surroundings less attractive to moose.

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Réhaume Courtois

Papers

Linking moose habitat selection to limiting factors

Effects of sampling effort based on GPS telemetry on home-range size estimations

Cumulative effects of forestry on habitat use by gray wolf (Canis lupus) in the boreal forest

Behavioural responses of moose to thermal conditions in the boreal forest

Behavior of Moose Relative to a Road Network