Allometric scaling of intraspecific space use

Allometric scaling relationships enable exploration of animal space-use patterns, yet interspecific studies cannot address many of the underlying mechanisms. We present the first intraspecific study of home range (HR) allometry relative to energetic requirements over several orders of magnitude of body mass, using as a model the predatory fish, pike Esox lucius. Analogous with interspecific studies, we show that space use increases more rapidly with mass (exponent = 1.08) than metabolic scaling theories predict. Our results support a theory that suggests increasing HR overlap with body mass explains many of these differences in allometric scaling of HR size. We conclude that, on a population scale, HR size and energetic requirement scale allometrically, but with different exponents.     

Landscape heterogeneity and the use of space by elephants in the Kruger National Park, South Africa

Landscape heterogeneity may influence ranging behaviour of mammals. Here we relate the home range size of elephants living in the Kruger National Park to the number of patches, proportion of each patch, spatial arrangement of patches, patch shape, and contrast between neighbouring patches. Home range sizes decreased exponentially with an increase in the number of patches per 100 km² and the home range sizes of bulls were in general more strongly related to measures of heterogeneity. This may reflect differences in perception of heterogeneity between the sexes.     

Individual space use strategies of wild juvenile Atlantic salmon

Movements of 60 stream-dwelling wild Atlantic salmon Salmo salar (97–118 mm), each tagged with a passive integrated transponder, were monitored during four trials in an enclosed section (24 m long, 45·1 m² total area) of a stream at a range of densities (four, eight, 16 and 32 fish per enclosure). Patterns of space use differed markedly between individuals, with 80% of fish establishing home ranges within 8 days of introduction to the enclosure (settlers) and the remainder continuing to move throughout the length of the enclosure (non-settlers). Although aggressive interactions were quite frequent and dominant fish were observed chasing subordinates, there was considerable overlap of home ranges of settlers at all densities; this was the case even at lower densities at which only a fraction of the enclosure was used by the fish. Thus, rather than adopting fixed territories, the salmon showed a high level of space sharing. Individual fish used the same local area in different ways, ranging from highly localized feeding on drifting food items to a wider-ranging strategy of specialising on benthic food. Among the fish that settled absolute growth rates were inversely related to body size, and at high densities fish lost weight. These results suggest that space use in wild juvenile salmon is more complex than a mosaic of territories, that salmon demonstrate significant variability in individual space use patterns, and that large fish may suffer disproportionately when populations are at the carrying capacity of their environment.     

Density‐dependent space use affects interpretation of camera trap detection rates

Camera traps (CTs) are an increasingly popular tool for wildlife survey and monitoring. Estimating relative abundance in unmarked species is often done using detection rate as an index of relative abundance, which assumes that detection rate has a positive linear relationship with true abundance. This assumption may be violated if movement behavior varies with density, but the degree to which movement behavior is density‐dependent across taxa is unclear. The potential confounding of population‐level relative abundance indices by movement would depend on how regularly, and by what magnitude, movement rate and home‐range size vary with density. We conducted a systematic review and meta‐analysis to quantify relationships between movement rate, home‐range size, and density, across terrestrial mammalian taxa. We then simulated animal movements and CT sampling to test the effect of contrasting movement scenarios on CT detection rate indices. Overall, movement rate and home‐range size were negatively correlated with density and positively correlated with one another. The strength of the relationships varied significantly between taxa and populations. In simulations, detection rates were related to true abundance but underestimated change, particularly for slower moving species with small home ranges. In situations where animal space use changes markedly with density, we estimate that up to thirty percent of a true change in relative abundance may be missed due to the confounding effect of movement, making trend estimation more difficult. The common assumption that movement remains constant across densities is therefore violated across a wide range of mammal species. When studying unmarked species using CT detection rates, researchers and managers should explicitly consider that such indices of relative abundance reflect both density and movement. Practitioners interpreting changes in camera detection rates should be aware that observed differences may be biased low relative to true changes in abundance. Further information on animal movement, or methods that do not depend on assumptions of density‐independent movement, may be required to make robust inferences on population trends.     

Fine‐scale movements and use of space by spotted hyaena (Crocuta crocuta) on Ongava Game Reserve,Namibia

We deployed Global Positioning System (GPS) collars on spotted hyaena, Crocuta crocuta, on Ongava Game Reserve in northern Namibia. We analysed the movement profiles recorded from three periods of fine temporal scale (15 min interval) GPS data – dry season data from a sub‐adult female (36 days) and a lactating adult female (54 days), and wet season data from the same adult female (55 days). The hyaenas both had similar daily activity patterns – at rest between 12.00 and 18.00 hours, with a peak of activity in the 2‐h period around sunrise. They exhibited one or two active periods each night, travelling up to 30 km and being active for up to 10 h. Daily rest sites were widely distributed across the reserve, typically located on elevated ground and never revisited on consecutive days. In the dry season, both hyaenas made extensive use of the reserve, plus adjacent areas in Etosha National Park (sub‐adult home range 240 km², adult home range 366 km²). The wet season data for the adult female showed a significant reduction of space used (home range 232 km²). However, their utility distributions showed a nonuniform use of space, with multiple areas of high‐density utilization located away from open terrain.     

Movement and habitat use by the marbled lungfish Protopterus aethiopicus Heckel 1851 in Lake Baringo, Kenya

The marbled lungfish, Protopterus aethiopicus, a recent introduction into Lake Baringo, Kenya is now an important commercial species there. Because little is known about its behaviour, we used ultrasonic telemetry to investigate its movements and use of habitat as part of a broader biological study. Twelve marbled lungfish were implanted with ultrasonic tags and tracked for variable periods between September 2001 and 2002. Two individuals were tracked for most of the study period. Daily movement ranged from little or none to 5.2 km. Mean hourly rates of movement for three fish located twice a day (morning and late afternoon) over several days suggested that individuals were active throughout the diel period. Maximum lake depth was about 3 m and fish utilized all depths greater than 1 m. Six home ranges described for four lungfish varied in size from 5.8 to 19.8 km ² and were occupied for between 2 and 4.5 months. Use of habitat and the movement of marbled lungfish in Lake Baringo appeared to be influenced more by biotic than abiotic factors.     

Nowhere to run: semi-permeable barriers affect pronghorn space use

Animal movement can mediate the ecological consequences of fragmentation; however, barriers such as fences, roads, and railways are becoming a pervasive threat to wildlife. Pronghorn (Antilocapra americana) habitat in western North America has been fragmented by roads, railways, and fences. Although pronghorn are sensitive to barriers, neither the relative permeability of different barriers to crossing nor their influence on space use have been quantified. We used a large global positioning system (GPS)-collar dataset of pronghorn (n = 1,010 animal-years) in Wyoming, USA, to first quantify the likelihood that pronghorn cross each of 5 different anthropogenic barriers, including fences, county roads, railroads, state highways, and interstate highways (i.e., interstates). Next, we assessed how each barrier influenced pronghorn space use during the winter as indexed by the area occupied, and daily displacement relative to the density of barriers on an individual's winter range. The semi-permeability of the 5 barriers varied substantially, with the interstate being the most severe barrier to pronghorn movement. Pronghorn were >300 times less likely to cross interstates compared to state highways. Although pronghorn space use was rarely influenced by barriers within individual core winter ranges, pronghorn space use was constrained by barriers on the buffered periphery of individual winter ranges. Despite their different permeability to movement, the density of fences and combined interstates and railroads had similarly negative effects on pronghorn space use. Our results illustrate that the degree to which pronghorn avoid crossing barriers may scale up to affect access to habitat. Additionally, our results indicate that the effects of barriers on habitat access are not proportional to their permeability. Our results add to a growing consensus that effective management of mobile species depends on understanding how different kinds of semi-permeable barriers influence access and use of habitats.     

Ecological and anthropogenic influences on space use by spotted hyaenas

Due to increasing human encroachment into the remaining habitat of many large carnivore species, there is an immediate need to understand the ecological and anthropogenic factors influencing carnivore space use decisions. In particular, knowledge of changes in space use in response to disturbance, and the costs associated with these changes, will be critical in guiding conservation efforts. To investigate the ecological factors influencing carnivore space use, we intensively radiotracked members of two large social groups (clans) of spotted hyaenas Crocuta crocuta in the Masai Mara National Reserve, Kenya. In addition, we studied the influence of livestock grazing by comparing space use between two study clans that differed dramatically in exposure to grazing. Logistic regression modeling indicated that space use in the absence of livestock was most influenced by the location of the clan's communal den. However, hyaenas were also found to select shrubland, areas of high prey density, and proximity to seasonal streams. Movements of hyaenas exposed to livestock grazing were most influenced by vegetation type, with a strong avoidance of open grass plains. Den location and prey density had less influence on space use decisions in the disturbed than the undisturbed clan. Livestock distribution did not directly influence hyaena movements either during daytime, when livestock were present, or at night. We suggest that direct livestock avoidance was unnecessary due to the observed increased use of vegetative cover by hyaenas exposed to grazing livestock. The greater distances from the den, and from areas of high prey density at which hyaenas were found in disturbed than undisturbed areas indicates potential energetic costs incurred by disturbed hyaenas. Our results therefore suggest that reduced vegetative cover, as is often found outside protected areas, may result in more dramatic modifications of hyaena movements in the presence of livestock.     

Step selection techniques uncover the environmental predictors of space use patterns in flocks of Amazonian birds

Understanding the behavioral decisions behind animal movement and space use patterns is a key challenge for behavioral ecology. Tools to quantify these patterns from movement and animal–habitat interactions are vital for transforming ecology into a predictive science. This is particularly important in environments undergoing rapid anthropogenic changes, such as the Amazon rainforest, where animals face novel landscapes. Insectivorous bird flocks are key elements of avian biodiversity in the Amazonian ecosystem. Therefore, disentangling and quantifying the drivers behind their movement and space use patterns is of great importance for Amazonian conservation. We use a step selection function (SSF) approach to uncover environmental drivers behind movement choices. This is used to construct a mechanistic model, from which we derive predicted utilization distributions (home ranges) of flocks. We show that movement decisions are significantly influenced by canopy height and topography, but depletion and renewal of resources do not appear to affect movement significantly. We quantify the magnitude of these effects and demonstrate that they are helpful for understanding various heterogeneous aspects of space use. We compare our results to recent analytic derivations of space use, demonstrating that the analytic approximation is only accurate when assuming that there is no persistence in the animals' movement. Our model can be translated into other environments or hypothetical scenarios, such as those given by proposed future anthropogenic actions, to make predictions of spatial patterns in bird flocks. Furthermore, our approach is quite general, so could potentially be used to understand the drivers of movement and spatial patterns for a wide variety of animal communities.     

Movement patterns of Cephalopholis cruentata in a marine reserve in St Lucia, W.I., obtained from ultrasonic telemetry

Home range size and shape, and within home range movement patterns were determined from 10 adult graysby Cephalopholis cruentata implanted with acoustic transmitters on two reefs within the Soufriere Marine Management Area, St Lucia. The mean home range area was 2120 m²(range 1200 to 4000 m²) and non‐circular in shape. All fish demonstrated a clear preference for a specific area in their home range where, on average, 72·9% of an individuals position were located. Time spent in this favoured area was correlated with reef complexity. Fish with greater activity and displacement possessed a larger home range. Activity and displacement were higher by night than by day, and consistent with this, fish spent less time in the preferred area of their home range by night than by day. Tagged individuals demonstrated a strong tendency to avoid sand habitat present at reserve boundaries.     

Does the presence of a human tracker affect the behaviour of radio‐tagged wood mice Apodemus sylvaticus?

1. While the effect on animals of handling them and fitting them with radio tags has been investigated, little work has been reported testing the effect of the presence of a human observer tracking an animal. 2. The activity of wood mice fitted with radio collars and confined to a semifield pen was measured in the presence and the absence of a human observer. Free‐ranging wood mice were radio tracked by using two protocols: ‘fixes’ taken periodically, and periods of continuous tracking. 3. There was no significant difference in any quantified aspect of the animals’ behaviour. This suggests that the presence of a human tracker has a negligible effect on the behaviour of wood mice.     

Influence of sea turtle nesting on hunting behavior and movements of jaguars in the dry forest of northwest Costa Rica

Jaguars (Panthera onca) are opportunistic predators that prey on large profitable prey items, such as sea turtles at nesting beaches. Here, we use jaguar and sea turtle track-count surveys, combined with satellite telemetry of one jaguar, to evaluate whether jaguar hunting behavior and movements are influenced by seasonal sea turtle nesting in the Sector Santa Rosa of Área de Conservación Guanacaste in northwest Costa Rica. We used generalized linear models to evaluate the effect of moon phase and sea surface temperature on olive ridley (Lepidochelis olivacea) and green turtle (Chelonia mydas) nesting abundance, as well as the combination of these predictors on the frequency of jaguar predation activity (proximity to nesting beaches) and movements. For home-range size and location analyses, we calculated kernel density estimates for each season at three different temporal scales. Sea turtle nesting season influenced jaguar activity patterns, as well as sea turtle abundance was related to jaguar locations and predation events, but jaguar home-range size (88.8 km² overall) did not differ between nesting seasons or among temporal scales. Environmental conditions influenced sea turtle nesting and, as a consequence, also influenced jaguar movements and foraging activity. Our study defined the home range of a female jaguar in the tropical dry forest and its relationship to seasonally abundant turtles. Additional information related to the effect of tourism on jaguar–sea turtle interactions would improve conservation of these species at unique nesting beaches in the area.     

Spatial ecology of jaguars,pumas, and ocelots: a review of the state of knowledge

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Wintering space use and site fidelity in a nomadic species,the snowy owl

Migratory species can exploit many habitats over vast geographic areas and adopt various patterns of space and habitat use throughout their annual cycle. In nomadic species, determinants of habitat use during the non‐breeding season are poorly known due to the unpredictability of their movement patterns. Here, we analysed variability in wintering space and habitat use by a highly nomadic species, the snowy owl, in eastern North America. Using 21 females tracked by satellite telemetry between 2007 and 2016, we 1) assessed how space use patterns in winter varied according to the type of environment (marine vs terrestrial), latitudinal zone (Arctic vs temperate), local snow conditions and lemming densities and 2) investigated winter habitat and site fidelity. Our results confirmed a high inter‐individual variation in patterns of habitat use by wintering snowy owls. Highly‐used areas were concentrated in the Arctic and in the marine and coastal environments. Owls wintering in the marine environment travelled over longer distances during the winter, had larger home ranges and these were divided in more smaller zones than individuals in terrestrial environments. Wintering home range sizes decreased with high winter lemming densities, use of the marine environment increased following high summer lemming densities, and a thick snow cover in autumn led to later settlement on the wintering ground. Contrary to expectations, snowy owls tended to make greater use of the marine environment when snow cover was thin. Snowy owls were highly consistent in their use of a given wintering environment and a specific latitudinal zone between years, but demonstrated flexibility in their space use and a modest site fidelity. The snowy owls’ consistency in wintering habitat use may provide them with advantages in terms of experience but their mobility and flexibility may help them to cope with changing environmental conditions at fine spatial scale.     

Staying close to home: Ecological constraints on space use and range fidelity in a mountain ungulate

Understanding patterns of animal space use and range fidelity has important implications for species and habitat conservation. For species that live in highly seasonal environments, such as mountain goats (Oreamnos americanus), spatial use patterns are expected to vary in relation to seasonal changes in environmental conditions and sex‐ or age‐specific selection pressures. To address hypotheses about sex, age, and seasonality influence on space‐use ecology, we collected GPS location data from 263 radio‐collared mountain goats (males, n = 140; females, n = 123) in coastal Alaska during 2005–2016. Location data were analyzed to derive seasonal and sex‐specific fixed‐kernel home range estimates and to quantify the degree of seasonal range and utilization distribution overlap. Overall, we determined that home range size was smallest during winter, expanded coincident with the onset of green‐up and parturition, and was largest during summer. Home range size of males and females did not differ significantly during winter, but females had larger home ranges than males during summer, a relationship that was switched during the mating season. Pairwise comparisons involving individual females across subsequent years indicated home ranges were significantly smaller during years when they gave birth to offspring. Mountain goats exhibited a strong degree of range fidelity, and 99% (n = 138) of individual animals returned to their previous year''s seasonal range with an average annual Bhattacharyya''s affinity utilization distribution overlap index of 68%. Similarity of seasonal home range utilization distributions varied in relation to sex and season in some respects. Home range overlap was highest during the summer vegetation growing season, particularly among females. These findings advance our understanding about how environmental variation and sex‐ and age‐related reproductive constraints influence space use and range fidelity among alpine ungulates. Documentation of the high degree of range fidelity among mountain goats has important conservation implications in landscapes increasingly altered by anthropogenic activities.     

Animal movement tools (amt): R package for managing tracking data and conducting habitat selection analyses

Advances in tracking technology have led to an exponential increase in animal location data, greatly enhancing our ability to address interesting questions in movement ecology, but also presenting new challenges related to data management and analysis. Step‐selection functions (SSFs) are commonly used to link environmental covariates to animal location data collected at fine temporal resolution. SSFs are estimated by comparing observed steps connecting successive animal locations to random steps, using a likelihood equivalent of a Cox proportional hazards model. By using common statistical distributions to model step length and turn angle distributions, and including habitat‐ and movement‐related covariates (functions of distances between points, angular deviations), it is possible to make inference regarding habitat selection and movement processes or to control one process while investigating the other. The fitted model can also be used to estimate utilization distributions and mechanistic home ranges. Here, we present the R package amt (animal movement tools) that allows users to fit SSFs to data and to simulate space use of animals from fitted models. The amt package also provides tools for managing telemetry data. Using fisher (Pekania pennanti) data as a case study, we illustrate a four‐step approach to the analysis of animal movement data, consisting of data management, exploratory data analysis, fitting of models, and simulating from fitted models.     

Home range and habitat use of translocated endangered species,Cottus koreanus,in South Korea

The home range and characteristics of Cottus koreanus were investigated using passive integrated transponder (PIT) telemetry in the Gulji Stream, Korea, where the target species was artificially translocated. After release, tagged individuals moved up to 78?m away from the release site in search of proper habitats. The average distance moved until settlement was 17.1?m. The observed home range of settled individuals had longitudinal sections of 9.9?±?3.6?m and surface areas of 7.2?±?2.7?m². This is comparable to congeneric species that inhabit similar ecological habitats. Once individuals had settled, they rarely moved from that site except during the spring season. The typical microhabitat characteristics of the sites where the released individuals settled are: water depth of 5–10?cm, water velocity of 0.1–0.3?m?s^?1, and the size of boulders and cobbles of 10–20?cm in diameter. This study of translocated C. koreanus individuals provides detailed information about habitats that can be used for effective habitat restoration and successful translocations attempts of this species.