Conservation of the Eurasian beaver Castor fiber: an olfactory perspective

• 1 Chemical communication in mammals includes an array of specific behaviours that are often ignored in terms of their potential relevance to conservation. Often used during territorial or social interactions between animals, chemical communication can also be used as a tool in reintroduction programmes. Reintroductions still exhibit high failure rates and methods to improve success should be investigated. The Eurasian beaver Castor fiber has been widely reintroduced across Europe after its near extinction in the 19th century.
• 2 Using olfactory studies in the beaver, we aim to demonstrate how scent transfers a range of information about the sender which can be used to monitor social and territorial behaviour along with general well‐being. Scent manipulation can be used to reduce human–beaver conflicts, and aid reintroduction success through reducing stress and territorial conflicts, and by influencing dispersal and settlement.
• 3 Two species of beavers, the Eurasian beaver and the North American beaver Castor canadensis, occupy freshwater habitats throughout North America and in parts of South America, most of Europe and parts of Asia. Most of the reviewed literature concerns the wild Eurasian beaver, its chemical communication and conservation; however, captive studies and those addressing North American beavers are also included.
• 4 Chemical communication is advanced and has been well documented in this highly territorial species. However, few studies directly link olfaction with conservation practices.
• 5 Olfactory studies in beavers can provide non‐invasive methods to monitor translocated animals and indicators of health. We conclude that chemical analysis, olfactory studies and behavioural manipulations involving semiochemicals have important impacts on conservation and can generate practical solutions to conservation problems including aiding animal capture, captive stress reduction, breeding pair formation and release site fidelity.

     

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.     

Time budgets and sex differences in the Eurasian beaver

The Eurasian beaver, Castor fiber, is one of the few obligate monogamous mammal species known, and nothing is known about the time budget of the mated pair. We investigated whether mated adult Eurasian beavers would display sex differences in time budgets. Using radiotracking, we obtained behavioural data on six mated pairs of adult beaver during 2000 and 2001 on two rivers in southeast Norway. Time budgets for males and females were compared in total and over the seasons, along with temperature data collected throughout the study. The three main behaviours of both males and females were travelling, foraging and being in the lodge, accounting for 92.0 and 93.2%, respectively, of each sex's overall time budgets. Time budgets did not differ between the sexes except that males allocated more time to travel. Time budgets for each sex did not vary with season, and water and air temperature data were positively correlated with time spent travelling for both sexes. These results support the hypothesis that male and female time budgets are similar, except for the time that males and females allocated to travel. We suggest that the similarities in behaviour result because parental care by both parents is essential to the successful rearing of offspring, leading to reduced behavioural sexual dimorphism; nevertheless, some differences still occur, perhaps relating to the male's indirect parental care. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Supplemental food alters nest defence and incubation behaviour of an open‐nesting wetland songbird

Climate‐driven increases in spring temperatures are expected to result in higher prey availability earlier in the breeding season for insectivorous birds breeding in wetland habitats. Predation during the incubation phase is a major cause of nesting failure in open‐nesting altricial birds such as the Eurasian reed warbler. The nest predation rate in this species has recently been shown to be substantially reduced under conditions of experimentally elevated invertebrate prey availability. Food availability near the nest may be an important determinant of adult incubation and nest defence behaviours during the incubation period. We used two experimental studies to compare incubation behaviour and nest defence in food‐supplemented and unsupplemented adult Eurasian reed warblers during the incubation phase. In the first study we measured nest defence behavioural responses to a taxidermic mount of a native predator (stoat Mustela erminea). In the second study we used temperature loggers installed in nests to measure breaks in incubation as a measure of nest vulnerability. Food‐supplemented birds responded aggressively to the presence of a predator more quickly than those in the unsupplemented group, suggesting they are closer to their nest and can more quickly detect a predator in the vicinity. Food‐supplemented birds also had shorter breaks in incubation (both in terms of maximum and mean off‐bout durations), presumably because they were foraging for shorter periods or over shorter distances from the nest. This study therefore identifies the behavioural mechanisms by which changes in food availability may lead to changes in nest survival and thus breeding productivity, in open‐nesting insectivorous birds.     

Biphasic Retention of One-Trial Learning in a Clonal Fish (<Emphasis Type="Italic">Poecilia formosa</Emphasis>)

Most species used for behavioural studies are bisexual. Sexual dimorphism determines genotypic diversity and behavioural variation within a species. The relative contribution of a genome to a specific behaviour is for the most part indiscernible, but gene changes can alter behaviour in many different ways¹. Within a species, strain differences can contribute to behavioural differences and many less clearly systematic behaviours, such as the aptitude to learn or to recall, may be genetically determined². Genotypic diversity, on the other hand, obscures gene correlates of behaviour because each organism brings a unique repertoire of behaviours to the experimental situation. Against this, learning research has found basic phenomena of learning and memory to be valid across many vertebrate species. In an effort to reduce genotypic and behavioural variability in studying memory processing, we have used a unisexual clonal fish, Poecilia formosa, as suggested by Agranoff and Davis³.     

Identifying spawning events of greater amberjack using accelerometers

Direct observations of spawning events of large free-swimming migratory fish are difficult in nature. However, behavioural changes specific to spawning events of fish species, i.e. swimming depth or acceleration of body motion using data loggers attached to the body, have successfully been used to identify spawning. In this study, we observed the spawning behaviour in an experimental tank of a pair of greater amberjack Seriola dumerili, which were matured by injection of gonadotropin, by attaching an acceleration data logger, an accelerometer. The male and female were observed to vibrate their bodies during the moment of spawning. Analysis of the acceleration data showed the simultaneous appearance of a dominant cycle (cycle indicating the highest amplitude in 1?s) under 0.4?s continuing over 7?sand dominant amplitude over 0.4 indicating the time of the spawning in both sexes. These characteristics are useful for the development of an algorithm to detect the spawning behaviour, allowing us to develop a new data logger with an on-board algorithm for detection of spawning behaviour. Before development of this system, we need to develop new algorithms for separation of spawning behaviour from other similar large motion behaviours, particularly from feeding and prey avoidance behaviour.     

Behavioural differences between two captive populations of red jungle fowl (Gallus gallus) with different genetic background,raised under identical conditions

Ex situ conservation of threatened species may lead to behavioural adaptation, which can affect success of reintroduction attempts. In previous studies, we investigated the effects of captivity on the behaviour of red jungle fowl (Gallus gallus) and found that captive populations differed behaviourally as well as genetically. The aim of the present study was to compare the behaviour of two of the previously studied populations, raised under identical conditions. Eggs were collected from birds at Copenhagen zoo (Cop) and Götala research station (Got) and were incubated and hatched together. Twenty-eight birds (16 Got and 12 Cop) were reared together and tested in eight different behavioural tests, measuring different aspects of fear-related behaviours as well as exploratory and social behaviours. The study revealed several differences in fear-related behaviours between the populations but none in exploratory or social behaviours. In general, one of the populations (Cop) showed more intense fear behaviours than the other (Got), which instead were less fearful in their behaviours. This indicates that breeding animals in captivity may lead to behavioural modifications, which can affect the outcome of reintroductions. The results further suggest that fear-related behaviours are dependent on the genetic background of the animals while social behaviours may be more influenced by the social environment. Since fear-related behaviours, such as predator avoidance and fear of humans, are essential for a life in the wild, these aspects are crucial for the breeding of animals in captivity for conservation purposes.     

A test of the dear enemy phenomenon in the Eurasian beaver

We tested the hypothesis that Eurasian beavers, Castor fiber, display the dear enemy phenomenon; that is, they respond less aggressively to intrusions by their territorial neighbours than to intrusions by nonterritorial floaters (strangers). This ability could be advantageous in facilitating differential treatment of wandering strangers versus established neighbours. Territorial beavers were presented with scent from neighbouring and stranger adult males. Thirty-nine different active beaver families, 18 in 1998 and 21 in 1999, were presented with a two-way choice between two pairs of experimental scent mounds; mounds with castoreum from a neighbour and a stranger, and mounds with anal gland secretion from a neighbour and a stranger. Direct observations of the families during evenings showed that: (1) beavers sniffed both castoreum and anal gland secretion from a stranger significantly longer than from a neighbour, and (2) beavers responded aggressively (stood on the mound on their hind feet, pawing and/or overmarking) significantly longer to castoreum, but not to anal gland secretion, from a stranger than from a neighbour. When experimental scent mounds were allowed to remain overnight and the beavers' responses were measured the following morning, the beavers' responses were significantly stronger to both castoreum and anal gland secretion from a stranger than from a neighbour. These findings indicate that Eurasian beavers can use scent to discriminate between neighbours and strangers, thereby supporting existence of the dear enemy phenomenon in this species. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Diving behavior in a free‐living,semi‐aquatic herbivore,the Eurasian beaver Castor fiber

Semi‐aquatic mammals have secondarily returned to the aquatic environment, although they spend a major part of their life operating in air. Moving both on land, as well as in, and under water is challenging because such species are considered to be imperfectly adapted to both environments. We deployed accelerometers combined with a depth sensor to study the diving behavior of 12 free‐living Eurasian beavers Castor fiber in southeast Norway between 2009 and 2011 to examine the extent to which beavers conformed with mass‐dependent dive capacities, expecting them to be poorer than wholly aquatic species. Dives were generally shallow (<1 m) and of short duration (<30 s), suggesting that the majority of dives were aerobic. Dive parameters such as maximum diving depth, dive duration, and bottom phase duration were related to the effort during different dive phases and the maximum depth reached. During the descent, mean vectorial dynamic body acceleration (VeDBA—a proxy for movement power) was highest near the surface, probably due to increased upthrust linked to fur‐ and lung‐associated air. Inconsistently though, mean VeDBA underwater was highest during the ascent when this air would be expected to help drive the animals back to the surface. Higher movement costs during ascents may arise from transporting materials up, the air bubbling out of the fur, and/or the animals’ exhaling during the bottom phase of the dive. In a manner similar to other homeotherms, beavers extended both dive and bottom phase durations with diving depth. Deeper dives tended to have a longer bottom phase, although its duration was shortened with increased VeDBA during the bottom phase. Water temperature did not affect diving behavior. Overall, the beavers’ dive profile (depth, duration) was similar to other semi‐aquatic freshwater divers. However, beavers dived for only 2.8% of their active time, presumably because they do not rely on diving for food acquisition.     

Landscape structure and population density affect intraspecific aggression in beavers

Intraspecific competition plays an important role for territory acquisition and occupancy, in turn affecting individual fitness. Thus, understanding the drivers of intraspecific aggression can increase our understanding of population dynamics. Here, we investigated intraspecific aggression in Eurasian (Castor fiber) and North American (Castor canadensis) beavers that are both monogamous, territorial mammals. Combined, we examined tail scars from >1,000 beavers (>2,000 capture events) as part of two long‐term studies in Norway and the USA. We investigated the influence of landscape structure, population density, sex, age, and (for Eurasian beavers only) social status and group size on the number of tail scars caused by conspecifics. The number of tail scars was affected by population density in well‐connected landscape types (large lakes and rivers), but not in more isolated areas (ponds), where individuals generally had fewer tail scars. Further, the relationship of population density was not linear. In the North American beaver population occurring in large lakes, intraspecific aggression increased with population density. Conversely, in the saturated Eurasian beaver population, intraspecific aggression was in a negative relationship with population density (except at the highest densities), likely due to inverse density‐dependent intruder pressure via dispersers. Our findings emphasize that population density can affect intraspecific aggression depending on landscape structure, which might have important consequences for local patterns of dispersal, mate change, and territory occupancy, all of which can affect population dynamics.     

More genetic data are needed before populations are mixed: response to ‘Sourcing Eurasian beaver Castor fiber stock for reintroductions in Great Britain and Western Europe’

Halley described three options for sourcing Eurasian beavers Castor fiber for reintroduction in Britain and Western Europe: (i) use animals from populations within a single western evolutionarily significant unit (ESU); (ii) mix animals from two or three western ESUs; and (iii) mix animals from eastern and western ESUs. Option three contravenes International Union for Conservation of Nature guidelines, so debate should focus on options one and two. We believe there is a need for further genetic analysis, before a final decision can be made. This decision should not be heavily influenced by cost or be determined by the genotypes of existing captive or escaped populations.     

Accelerometry predicts daily energy expenditure in a bird with high activity levels

Animal ecology is shaped by energy costs, yet it is difficult to measure fine-scale energy expenditure in the wild. Because metabolism is often closely correlated with mechanical work, accelerometers have the potential to provide detailed information on energy expenditure of wild animals over fine temporal scales. Nonetheless, accelerometry needs to be validated on wild animals, especially across different locomotory modes. We merged data collected on 20 thick-billed murres (Uria lomvia) from miniature accelerometers with measurements of daily energy expenditure over 24 h using doubly labelled water. Across three different locomotory modes (swimming, flying and movement on land), dynamic body acceleration was a good predictor of daily energy expenditure as measured independently by doubly labelled water (R² = 0.73). The most parsimonious model suggested that different equations were needed to predict energy expenditure from accelerometry for flying than for surface swimming or activity on land (R² = 0.81). Our results demonstrate that accelerometers can provide an accurate integrated measure of energy expenditure in wild animals using many different locomotory modes.     

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 variations of the digestive tract of the Eurasian beaver <Emphasis Type="Italic">Castor fiber</Emphasis>

Forage availability for wild rodents varies with season. In turn, the composition of food can affect morphometric parameters of the digestive tract. This study was performed in Eurasian beavers (Castor fiber) whose population was close to extinction in most Eurasian countries, but has now increased. Due to the previous low number of studies, information about the effect of forage availability on the digestive tract morphology has previously been lacking. This study was performed using beavers captured from the natural environment during three seasons of different forage availability: winter, summer and autumn. It was found that the diet of the beaver varied during the year; in winter it was dominated by woody material consisting of willow shoots, whereas in summer the diet was primarily herbs, grass and leaves. Season also affected the mass of digested contents of the digestive tract. The digestive content increased in the caecum and colon in winter and autumn, when poor-quality food dominated the beaver’s diet. The results indicated that the digestive tract parameters of beavers varied based on the composition of available forage.     

Behavioural effects of temperature on ectothermic animals: unifying thermal physiology and behavioural plasticity

Temperature imposes significant constraints on ectothermic animals, and these organisms have evolved numerous adaptations to respond to these constraints. While the impacts of temperature on the physiology of ectotherms have been extensively studied, there are currently no frameworks available that outline the multiple and often simultaneous pathways by which temperature can affect behaviour. Drawing from the literature on insects, we propose a unified framework that should apply to all ectothermic animals, generalizing temperature's behavioural effects into: (1) kinetic effects, resulting from temperature's bottom‐up constraining influence on metabolism and neurophysiology over a range of timescales (from short to long term), and (2) integrated effects, where the top‐down integration of thermal information intentionally initiates or modifies a behaviour (behavioural thermoregulation, thermal orientation, thermosensory behavioural adjustments). We discuss the difficulty in distinguishing adaptive behavioural changes from constraints when observing animals' behavioural responses to temperature. We then propose two complementary approaches to distinguish adaptations from constraints, and categorize behaviours according to our framework: (i) ‘kinetic null modelling’ of temperature's effects on behaviour; and (ii) behavioural ecology experiments using temperature‐insensitive mutants. Our framework should help to guide future research on the complex relationship between temperature and behaviour in ectothermic animals.     

Ontogenetic intraspecific allometry of autochthonous and reintroduced Eurasian beaver <Emphasis Type="Italic">Castor fiber</Emphasis> populations

Ontogenetic allometry of the cranium was studied in 580 individuals of the Eurasian beavers Castor fiber in two subspecies out of six autochthonous and reintroduced populations using the methods of multivariate nonparametric statistics. It was noted that the growth dynamics demonstrates a correspondence to trajectories of ontogenetic rodent models with the difference that intensive beaver growth finishes by the age of four years. Significant differences in the dynamics of age variability were detected in the beavers of two subspecies; with somewhat smaller initial postnatal sizes, C. fiber belorussicus grows faster than C. fiber orientoeuropaeus.     

Stereotypic route-tracing in experimentally caged songbirds correlates with general behavioural disinhibition

Repetitive, unvarying and apparently functionless behaviours called stereotypies are common in caged animals, but the mechanisms of cage stereotypy have remained elusive. We found that stereotypies correlate with a sign of altered brain functioning, the general disinhibition of behaviour, found in stereotyping human patients and animals treated with psychostimulants. We investigated route-tracing stereotypy in blue tits, Parus caeruleus, and marsh tits,P. palustris , caged in a behavioural research laboratory. In experiment 1, stereotypy correlated with disinhibition of responses, shown as persistent responding in extinction learning. In experiment 2a, stereotypy in blue tits correlated with repetitive sequence generation on a ‘gambling task’ that specifically measures disinhibited striatal functioning in humans and correlates with stereotypy in autistic and schizophrenic patients. In experiment 2b, the behavioural disinhibition correlated with stereotypy also affected food-storing behaviour in marsh tits, in particular their response to cache pilfering. In experiment 3, the sequencing of nonstereotypic home cage behaviour was correlated with stereotypy. Finally, changes in stereotypy induced by environmental enrichment correlated with changes in these measures of altered behavioural control. These results suggest that housing conditions that cause stereotypy thus alter many aspects of the behavioural control of caged subjects, mediated by altered striatal functioning. The implications for understanding cage stereotypies in laboratory, farm and zoo animals, and for laboratory-based behavioural experiments, are discussed. We suggest that improving housing conditions so that cage stereotypies do not develop would enhance the validity of laboratory-based behavioural research.     

Haematology and Serum Biochemistry Parameters and Variations in the Eurasian Beaver (Castor fiber)

Haematology parameters (N = 24) and serum biochemistry parameters (N = 35) were determined for wild Eurasian beavers (Castor fiber), between 6 months – 12 years old. Of the population tested in this study, N = 18 Eurasian beavers were from Norway and N = 17 originating from Bavaria but now living extensively in a reserve in England. All blood samples were collected from beavers via the ventral tail vein. All beavers were chemically restrained using inhalant isoflurane in 100% oxygen prior to blood sampling. Results were determined for haematological and serum biochemical parameters for the species and were compared between the two different populations with differences in means estimated and significant differences being noted. Standard blood parameters for the Eurasian beaver were determined and their ranges characterised using percentiles. Whilst the majority of blood parameters between the two populations showed no significant variation, haemoglobin, packed cell volume, mean cell haemoglobin and white blood cell counts showed significantly greater values (p<0.01) in the Bavarian origin population than the Norwegian; neutrophil counts, alpha 2 globulins, cholesterol, sodium: potassium ratios and phosphorus levels showed significantly (p<0.05) greater values in Bavarian versus Norwegian; and potassium, bile acids, gamma globulins, urea, creatinine and total calcium values levels showed significantly (p<0.05) greater values in Norwegian versus Bavarian relict populations. No significant differences were noted between male and female beavers or between sexually immature (<3 years old) and sexually mature (≥3 years old) beavers in the animals sampled. With Eurasian beaver reintroduction encouraged by legislation throughout Europe, knowledge of baseline blood values for the species and any variations therein is essential when assessing their health and welfare and the success or failure of any reintroduction program. This is the first study to produce base-line blood values and their variations for the Eurasian beaver.     

Effects of fleas on nest success of Arctic barnacle geese: experimentally testing the mechanism

Parasites have detrimental effects on their hosts’ fitness. Therefore, behavioural adaptations have evolved to avoid parasites or, when an individual is already in contact with a parasite, prevent or minimize infections. Such anti‐parasite behaviours can be very effective, but can also be costly for the host. Specifically, ectoparasites can elicit strong host anti‐parasite behaviours and interactions between fleas (Siphonaptera) and their hosts are one of the best studied. In altricial bird species, nest fleas can negatively affect both parent and offspring fitness components. However, knowledge on the effects of fleas on precocial bird species is scarce. Research on geese in the Canadian Arctic indicated that fleas have a negative impact on reproductive success. One possible hypothesis is that fleas may affect female incubation behaviour. Breeding females with many fleas in their nest may increase the frequency and/or duration of incubation breaks and could even totally desert their nest. The aim of our study was to 1) determine if a similar negative relationship existed between flea abundance and reproductive success in our study colony of Arctic breeding barnacle geese Branta leucopsis and 2) experimentally quantify if such effects could be explained by a negative effect of nest fleas on female behaviour. We compared host anti‐parasite and incubation behaviour between experimentally flea‐reduced and control nests using wildlife cameras and temperature loggers. We found that flea abundance was negatively associated with hatching success. We found little experimental support, however, for changes in behaviour of the breeding female as a possible mechanism to explain this effect.     

Extending the Functionality of Behavioural Change-Point Analysis with k-Means Clustering: A Case Study with the Little Penguin (Eudyptula minor)

We present a simple framework for classifying mutually exclusive behavioural states within the geospatial lifelines of animals. This method involves use of three sequentially applied statistical procedures: (1) behavioural change point analysis to partition movement trajectories into discrete bouts of same-state behaviours, based on abrupt changes in the spatio-temporal autocorrelation structure of movement parameters; (2) hierarchical multivariate cluster analysis to determine the number of different behavioural states; and (3) k-means clustering to classify inferred bouts of same-state location observations into behavioural modes. We demonstrate application of the method by analysing synthetic trajectories of known ‘artificial behaviours’ comprised of different correlated random walks, as well as real foraging trajectories of little penguins (Eudyptula minor) obtained by global-positioning-system telemetry. Our results show that the modelling procedure correctly classified 92.5% of all individual location observations in the synthetic trajectories, demonstrating reasonable ability to successfully discriminate behavioural modes. Most individual little penguins were found to exhibit three unique behavioural states (resting, commuting/active searching, area-restricted foraging), with variation in the timing and locations of observations apparently related to ambient light, bathymetry, and proximity to coastlines and river mouths. Addition of k-means clustering extends the utility of behavioural change point analysis, by providing a simple means through which the behaviours inferred for the location observations comprising individual movement trajectories can be objectively classified.