Evaluation of attractant flavours for use in oral vaccine baits for badgers (<Emphasis Type="Italic">Meles meles</Emphasis>)

European badgers (Meles meles) are a wildlife reservoir for Mycobacterium bovis infection (tuberculosis) in Ireland and the UK and are implicated in the transmission of infection to livestock. Vaccination of badgers with the human BCG vaccine (Bacille Calmette Guerin) is considered as an important strategy to reduce the burden of disease in this species, and a pragmatic approach is likely to involve oral vaccination. In this study, we evaluated nine different flavours for use as attractants in a prototype oral vaccine bait for European badgers (M. meles): aniseed, apple, cocoa powder, carob powder, curry, fish, garlic, peanut and strawberry. The bait matrix was composed of a natural lipid formulation, developed as a vehicle for oral vaccination against tuberculosis in wildlife. A ‘food for work’ paradigm was employed during the trials to ensure the animals were actively seeking the baits. The trials showed carob and cocoa powders were equally attractive and more attractive than any of the other candidates. Carob and cocoa show potential as bait attractants for badgers and might form part of a novel vaccine delivery system.     

How to reliably estimate the tortuosity of an animal's path: straightness, sinuosity, or fractal dimension?

The tortuosity of an animal's path is a key parameter in orientation and searching behaviours. The tortuosity of an oriented path is inversely related to the efficiency of the orientation mechanism involved, the best mechanism being assumed to allow the animal to reach its goal along a straight line movement. The tortuosity of a random search path controls the local searching intensity, allowing the animal to adjust its search effort to the local profitability of the environment. This paper shows that (1) the efficiency of an oriented path can be reliably estimated by a straightness index computed as the ratio between the distance from the starting point to the goal and the path length travelled to reach the goal, but such a simple index, ranging between 0 and 1, cannot be applied to random search paths; (2) the tortuosity of a random search path, ranging between straight line movement and Brownian motion, can be reliably estimated by a sinuosity index which combines the mean cosine of changes of direction with the mean step length; and (3) in the current state of the art, the fractal analysis of animals' paths, which may appear as an alternative and promising way to measure the tortuosity of a random search path as a fractal dimension ranging between 1 (straight line movement) and 2 (Brownian motion), is only liable to generate artifactual results. This paper also provides some help for distinguishing between oriented and random search paths, and depicts a general, comprehensive framework for analysing individual animals' paths in a two-dimensional space.     

Animal movement in dynamic landscapes: interaction between behavioural strategies and resource distributions

Most ecological and evolutionary processes are thought to critically depend on dispersal and individual movement but there is little empirical information on the movement strategies used by animals to find resources. In particular, it is unclear whether behavioural variation exists at all scales, or whether behavioural decisions are primarily made at small spatial scales and thus broad-scale patterns of movement simply reflect underlying resource distributions. We evaluated animal movement responses to variable resource distributions using the grey teal (Anas gracilis) in agricultural and desert landscapes in Australia as a model system. Birds in the two landscapes differed in the fractal dimension of their movement paths, with teal in the desert landscape moving less tortuously overall than their counterparts in the agricultural landscape. However, the most striking result was the high levels of individual variability in movement strategies, with different animals exhibiting different responses to the same resources. Teal in the agricultural basin moved with both high and low tortuosity, while teal in the desert basin primarily moved using low levels of tortuosity. These results call into question the idea that broad-scale movement patterns simply reflect underlying resource distributions, and suggest that movement responses in some animals may be behaviourally complex regardless of the spatial scale over which movement occurs.     

Badger <Emphasis Type="Italic">Meles meles</Emphasis> feeding ecology in dry Mediterranean environments of the southwest edge of its distribution range

The dietary preference of badgers Meles meles (Linnaeus, 1758) for earthworms and other food resources has been widely discussed. In the Mediterranean area, rabbits Oryctolagus cuniculus, fruits (mainly olives Olea europaea) and arthropods are the most important foods. We describe badger dietary habits in a xeric Mediterranean area, comparing use and availability of olives and rabbits. Our results indicated that olives and rabbits are the predominant resources consumed, however we did not detect any evidence of specialisation when we compared consumption versus field availability. In the case of olives we found that their consumption is reduced when juvenile rabbits or figs Ficus carica are widely available, which is evidence against any specialisation in this resource. Figs appear to be a key food item for badgers; they were consumed in large amounts when available. Badgers in this xeric area can be viewed as generalist or facultative specialists, using the most profitable resource when available but shifting its preferences to other less profitable food resources when availability of other primary food resource are reduced. Our results also indicated the high suitability of some human agricultural uses (eg fruit orchards) for this species, especially in the otherwise harsh Mediterranean environments.     

The effects of artificial and natural barriers on the movement of small mammals in Banff National Park, Canada

Barriers that dissect continuous habitat, such as roads, initiate the process of habitat fragmentation and can filter or eliminate animal movement through otherwise pristine areas. A severe road barrier appears to exist in Banff National Park where the Trans‐Canada Highway (TCH) is known to impede the movement of some animals, but the effects of this barrier relative to natural barriers and continuous habitat have not been assessed. To compare animal movement in response to these three contexts, we translocated three species of murid rodents (meadow voles, Microtus pennsylvanicus, deer mice, Peromyscus maniculatus, and red‐backed voles, Clethrionomys gapperi) across the TCH (an artificial barrier), a forested strip in the median of the TCH (a natural barrier), in the adjacent highway verge (continuous grassy) and in nearby forest (continuous forested). We coated murids with fluorescent dye, released them at standardized distances, and followed the resulting trails to obtain detailed information on movement paths. Overall, individuals were 20% less successful crossing the TCH than natural (forested) barriers and 10% less successful crossing natural barriers than continuous habitat, but exhibited marked variation among species. Path tortuosity was negatively related to return success; more convoluted paths were associated with lower success across all treatments. Deer mice returned to their home‐ranges more often than the other two species, perhaps because of their generalist habitat preferences, nocturnal activities, or larger home ranges. In all treatments, the fractal dimensions of movement paths were very qualitatively similar to those observed for return success and tortuosity, indicating that the responses of these species were not influenced by spatial scale. Together, our results suggest that (a) murid species differ in the movement limitation imposed by both natural and artificial barriers, perhaps as a consequence of differences in habitat preferences and activity patterns, and (b) murid responses to a road barrier are only moderately different from responses to the natural barrier of a forest edge in the context of translocation experiments.     

Colony and Individual Forager Responses to Food Quality in the New World Bumble Bee, Bombus occidentalis

The ability of a successful forager to activate colony foraging allows colonies to rapidly exploit ephemeral resources and is an important innovation in the evolution of sociality. We tested the ability of the species, Bombus occidentalis, to stimulate colony foraging for food varying in quality. We then analyzed the behavior of successful foragers inside the nest to learn more about potential foraging activation movements. The number of bees entering a foraging arena was positively correlated with food sucrose concentration (0.5, 1.0, and 2.5 M sucrose, equal to 16–65% w/w). Foragers spent significantly more time imbibing higher concentration solutions. Foragers then returned to the nest where they moved in elaborate paths at variable speeds. There was no significant effect of sucrose concentration on average forager velocity or time spent inside the nest. However, the length of a forager’s path inside the nest (total of all distances moved each 0.1 s) significantly increased with sucrose concentration. On average, individuals foraging on 2.5 M and 1.0 M solution walked paths respectively 1.6 fold and 1.4 longer than the paths of individuals foraging on 0.5 M solution. These longer paths could result in a greater number of nestmate contacts, a factor shown to be important in the activation of B. impatiens foragers and also reported in B. terrestris foragers.     

Turn designation, sampling rate and the misidentification of power laws in movement path data using maximum likelihood estimates

Many authors have claimed to observe animal movement paths that appear to be Lévy walks, i.e. a random walk where the distribution of move lengths follows an inverse power law. A Lévy walk is known to be the optimal search strategy of a particular class of random walks in certain environments; hence, it is important to distinguish correctly between Lévy walks and other types of random walks in observed animal movement paths. Evidence of a power law distribution in the step length distribution of observed animal movement paths is often used to classify a particular movement path as a Lévy walk. However, there is some doubt about the accuracy of early studies that apparently found Lévy walk behaviour. A recently accepted method to determine whether a movement path truly exhibits Lévy walk behaviour is based on an analysis of move lengths with a maximum likelihood estimate using Akaike weights. Here, we show that simulated (non-Lévy) random walks representing different types of animal movement behaviour (a composite correlated random walk; pooled data from a set of random walks with different levels of correlation and three-dimensional correlated random walks projected into one dimension) can all show apparent power law behaviour typical of Lévy walks when using the maximum likelihood estimation method. The probability of the movement path being identified as having a power law step distribution is related to both the sampling rate used by the observer and the way that ‘turns’ or ‘reorientations’ in the movement path are designated. However, identification is also dependent on the nature and properties of the simulated path, and there is currently no standard method of observation and analysis that is robust for all cases. Our results indicate that even apparently robust maximum likelihood methods can lead to a mismatch between pattern and process, as paths arising from non-Lévy walks exhibit Lévy-like patterns.     

Intraspecific,size-dependent variation in the movement behaviour of a brackish-water isopod: a resource-free laboratory experiment

Intraspecific variability represents an important, yet inadequately investigated factor affecting the movement behaviour and ecology of mobile organisms. Here, the influence of sex, seasonality and body size on the movement behaviour of the brackish isopod Lekanesphaera hookeri were examined under resource-free laboratory conditions. The mean step length, total path length and average speed were determined twice during the year for adult and juvenile isopods encompassing a 10-fold range in body length. The scale-independent fractal dimension D was used to quantify the tortuosity of the movement paths. No relationships were observed between sex or season and all the movement metrics. In contrast, isopods' body size scaled negatively with the fractal dimension D of movement paths with a breakpoint at 2.6 mm, roughly corresponding to the size of morphometric maturation. No other relationships were observed between the body length of isopods and mean step length, total path length and average speed. The results indicate a sex- and season-independent ontogenetic shift in movement behaviour in L. hookeri. This suggests that in sphaeromatid isopods post-embryonic development determines not only continuous variations in size and proportions, but also a discontinuous change in the movement strategy adopted to interact with the surrounding space. Overall, these findings underscore the need to account explicitly for such changes in models predicting the spatial distribution of organisms characterised by wide intra-population size variation.     

Mantled Howler (<Emphasis Type="Italic">Alouatta palliata</Emphasis>) Arboreal Pathway Networks: Relative Impacts of Resource Availability and Forest Structure

Numerous studies have concluded that primates move about their environments in a nonrandom manner, frequently traveling between consecutive foraging sites along relatively straight-line paths. However, primates do not always take the most direct path between resources, and a number of species have been observed to travel repeatedly along a network of the same arboreal pathways. In this study, I used spatially explicit techniques to examine quantitatively what mantled howler monkey groups on Barro Colorado Island, Panama, accomplish by selecting nonlinear paths between resources and by repeatedly using the same paths within an arboreal network. Results show that chosen arboreal paths between sites where foraging occurred have higher levels of resource availability and canopy connectivity than comparable straight-line paths between the same sites. When comparing the relative importance of these factors, autologistic models of pathway choice indicate that though canopy connectivity is related to the location of repeatedly used arboreal pathway networks, the most statistically significant predictor is resource availability (both on a path and within a visual detection distance of a path). These results provide support for the hypothesis that repeated use of arboreal pathway networks aids in resource monitoring and acquisition. In addition, statistical models developed from 1 primary focal group’s travel patterns had high predictive value when employed to generate likely locations for arboreal pathways in the home ranges of 3 neighboring groups. This finding has important implications for studies of primate habitat use and seed dispersal, as it suggests that different groups consistently use similar characteristics when deciding on travel paths.     

Annual dynamics of den use by red foxesVulpes vulpes and badgersMeles meles in central Poland

This study on den use by red foxesVulpes vulpes (Linnaeus, 1758) and badgersMeles meles (Linnaeus, 1758) was conducted in central Poland (51°48’N, 19°53’E) over two years. We found a strong negative correlation between monthly den utilisation rate by foxes and air temperature (r=−0.83,p=0.0001). Correlation between temperature and den use by badgers was positive but less distinct (r=0.40,p=0.036). During the whole study period, a negative correlation between the monthly intensity of den use by badgers and by foxes was found (r=−0.51,p=0.008). The seasonal pattern of den utilisation differed between the species. Foxes used the dens mainly in winter and spring whereas in summer and autumn the intensity of den utilisation sharply decreased to a minimum in August-September. A peak in den use by foxes was observed in March and coincided with rut and selection of future breeding dens. In badgers, two peaks in den use were observed: the first in April and the second in August-September. An individual badger used more dens and used them more evenly throughout the year than an individual fox. The same dens were visited by foxes and badgers mostly from January to April, with a maximum in March.     

Negative or positive density‐dependence in movements depends on climatic seasons: The case of a Neotropical marsupial

One of the major challenges in animal ecology is to understand the factors and processes driving movement behaviour. Although density may influence movement patterns, the occurrence and nature of density‐dependence in animal movements are still unclear, particularly whether it may vary among populations of a species, or across time within a population. Here, we evaluate the occurrence and nature of density‐dependence in the movements of a Neotropical marsupial, the Grey four‐eyed opossum Philander frenatus (Didelphidae, Didelphimorphia). We quantified fine‐scale path tortuosity of individuals inhabiting continuous forest areas and forest fragments, in different climatic seasons (humid vs. super‐humid). We also determined the relative importance of population size compared to sex and body mass on movements, using a model‐selection approach. In forest fragments, path tortuosity increased with population size in the super‐humid season, but decreased in the humid season. In the continuous forest, path tortuosity was affected only by sex and body mass, being slightly higher in males and negatively related to body mass. The occurrence of density‐dependence on movements only in forest fragments is likely to reflect the higher overall density of P. frenatus in small forest fragments. The variation in the nature of density‐dependence between climatic seasons is likely to reflect a trade off between foraging over large areas (humid season, low resource availability) versus avoiding agonistic encounters (super‐humid season, high resource availability). Our results show that (i) density‐dependence in movements may be context‐dependent occurring only in areas of relatively high overall population density; and (ii) density may affect movements in different ways at different climatic seasons.     

Spatial movement optimization in Amazonian <Emphasis Type="Italic">Eciton burchellii</Emphasis> army ants

Foraging army ants face a problem general to many animals—how best to confront resource depletion and environmental heterogeneity. Army ants have presumably evolved a nomadic lifestyle as a way to minimize re-exploitation of previously foraged areas. However, this solution creates a challenge for an army ant colony: foraging by this colony and others creates a shifting landscape of food resources, where colonies should theoretically avoid their own previous foraging paths as well as those of other colonies. Here, we examine how colonies exploit this resource mosaic, using some of the optimality arguments first proposed and tested by Franks and Fletcher (1983), but with much larger data sets in a new location in SW Amazonia. Our data supported Franks and Fletcher’s (1983) model for systematic avoidance of raided areas during the statary phase, as well as a hypothesis of distance optimization between successive statary bivouacs. We also test and find significant evidence that foraging raids turn in opposite directions from the previous day’s directional angles more frequently than what would be expected if turning angles were distributed at random, which acts to move a colony away from recently exploited areas. This implies that colonies follow a straighter line path during the nomadic phase as opposed to a curved one, which acts to maximize distance between statary bivouacs. In addition to intra-colony movement optimization, we examine evidence for inter-colony avoidance from more than 330 colony emigrations and suggest that colony-specific pheromones are not necessarily repulsive to other colonies. Lastly, we compare our results with those of similar studies carried out at Barro Colorado Island (BCI), Panama. Despite a higher density of army ants in the SW Amazon region, colonies spend less time emigrating than their counterparts at BCI, which suggests a higher prey density in SW Amazonia.     

Isolating structures of gerbils’ digestive tract (Gerbillidae, <Emphasis Type="Italic">Rhombomys,Meriones</Emphasis>) and their functional significance

The morphology of the digestive tract of three species of gerbils characterized by different food specialization—Rhombomys opimus, Meriones tamariscinus and M. meridianus—is considered. The correlation between areas of glandular and cornified epithelium is variable and depends greatly on the type of stomach filling and distribution of food mass in it. The bordering fold and ileocecal junction is not only able to isolate the forestomach and caecum significantly but also may be a mechanism of regulation of food movement through the digestive tract. The close location of ileum’s opening to the caecum and its outlet to the colon, as well as isolation of caecum’s cavity by a circular fold, could provide direct food transit from the ileum to the colon.     

Spatio-temporal ecology and density of badgers <Emphasis Type="Italic">Meles meles</Emphasis> in the Swiss Jura Mountains

This study reports for the first time data on the spatio-temporal ecology of badgers living in a cold and wet mountain region (Swiss Jura Mountains). The home range, movements, activity patterns and habitat use of three badgers (two males, one female) were examined using radiotelemetry. Average home range size was 320 ha (MCP 100%), but the ranging behaviour of badgers varied at a seasonal scale. As in other regions, badgers were strictly nocturnal or crepuscular and showed a marked reduction of activity in the winter period. From spring to autumn, animals were active for an average (±SD) of 8.1 ± 2.4 h and travelled up to 9,460 m each night (mean±SD, 5,160 ± 2,600 m). The nightly distance travelled by badgers was positively correlated with their travel speed, the duration of the activity period and the used area, but not with night length. Year-round, the radio-collared animals avoided pastures and the vicinity of houses during their night trips. In winter and spring, individual badgers used forests and wooded pastures more than expected according to their availability, whereas cereal fields were actively selected in summer and autumn. Den-watching, night-lighting and radio-tracking data suggest that badgers live in pairs in this wet and cold region. Population density estimates range from 0.4 to 1.5 individuals/100 ha. We discuss the importance of trophic resources and climate as factors influencing badger behavioural ecology.     

Effect of Starvation on the Searching Path of the Predatory Bug <Emphasis Type="Italic">Deraeocoris Lutescens</Emphasis>

As physiological factors may influence the predatory behavior of insects, this study was designed to evaluate the possible impact that hunger has on the search path of a polyphagous predator: Deraeocoris lutescens Schilling (Heteroptera: Miridae). D. lutescens was fed a diet of Rhopalosiphum padi (Linné) (Homoptera: Aphididae) nymphs and Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) eggs at 21±1 °C and a photoperiod of 16:8 (L:D). After moulting, third and fifth instar nymphs and adults were placed in Petri dishes containing an abundant food source for 24 h. After 24 h, the food was removed and groups were randomly assigned to increasing periods of prey deprivation. Each D. lutescens nymph or adult was placed at the center of an empty arena and its searching path was recorded to calculate different path parameters and to determine the type of movement mode adopted. Results show an effect of starvation on both nymphal and adult D. lutescens searching paths. Satiated third instar nymphs manifested an intensive search path whereas fifth instar nymphs and adults adopted an intermediate one. When starved, the extensive search path appeared early in young nymphs, and later in the last instar nymphs and adults, according to the degree of prey deprivation.     

Anisotropy and shift of search behavior in Malabar grouper (<Emphasis Type="Italic">Epinephelus malabaricus</Emphasis>) larvae in response to prey availability

In order to enhance encounters with prey, planktonic predators may display different swimming behavior with respect to food availability and distribution. In this study, we used 3D video techniques to record the swimming behavior of malabar grouper (Epinephelus malabaricus) larvae in both the absence and the presence of prey (Artemia sp. nauplii). Swimming properties were investigated in all of the 3D, the two vertical, and the horizontal projections using scale-dependent (mean speed and Net to Gross Displacement Ratio) and scale-independent (fractal dimension) metrics. When prey was added, larvae swam slower and in a less convoluted way as compared to what was observed in the absence of food. The results obtained with scale-dependant metrics were confirmed by those obtained with scale-independent analyses. Both unveiled the anisotropy of the swimming behavior of grouper larvae that tend to swim toward the vertical axis in order to maximize encounters with prey patches. This study shows that malabar grouper larvae can optimize their search volume by switching their behavior and further draws attention to the need to consider both vertical- and horizontal-projections components while addressing the plankter’s swimming trajectories.     

Combining animal movements and behavioural data to detect behavioural states

Animal movement paths show variation in space caused by qualitative shifts in behaviours. I present a method that (1) uses both movement path data and ancillary sensor data to detect natural breakpoints in animal behaviour and (2) groups these segments into different behavioural states. The method can also combine analyses of different path segments or paths from different individuals. It does not assume any underlying movement mechanism. I give an example with simulated data. I also show the effects of random variation, # of states and # of segments on this method. I present a case study of a fisher movement path spanning 8 days, which shows four distinct behavioural states divided into 28 path segments when only turning angles and speed were considered. When accelerometer data were added, the analysis shows seven distinct behavioural states divided into 41 path segments.     

Restricted ranging behaviour in a high-density population of urban badgers

By studying the responses of different species to urbanization, it is possible to understand the impact of this type of habitat modification and to explore, more generally, the link between variations in the environment and changes in behaviour. We radio collared 17 badgers Meles meles from six social groups in a 1 km² urban study area in Brighton, UK, where local badger population density was high, and collected data on their ranging behaviour between 2005 and 2007. We aimed to determine how badgers adapt their behaviour to an urban environment and to assess the generality of previously reported differences in the ranging and territorial behaviour of urban and rural badgers. Analysis of habitat preferences and movement patterns suggested that garden habitat was principally used for foraging, while scrub and allotment habitats were important in allowing animals to travel from one part of their range to another. Group and individual home ranges were the smallest so far recorded for badgers (mean 100% minimum convex polygons=9.26 and 4.91 ha, respectively). Individual range size was negatively correlated with the availability of garden habitat, suggesting that the rich food resources provided by gardens enabled ranges to be small. Group ranges were mostly non-contiguous and there was no evidence of territorial scent marking; rather, activity was mainly restricted to areas in the vicinity of main setts. It is clear that badgers can adapt successfully to urban habitats and that this process affects various aspects of their behaviour. However, our high-density population of urban badgers displayed patterns of behaviour that differed not only from those of typical rural badgers, but also in some respects from those of a previously studied low-density urban population. We conclude that generalizations about the effects of urbanization must be made with caution.     

Predicting the Competitive Regime of Female <Emphasis Type="Italic">Colobus vellerosus</Emphasis> from the Distribution of Food Resources

We examined the spatial and temporal distribution of the foods of ursine colobus (Colobus vellerosus) at Boabeng-Fiema, Ghana as a means to predict the monopolizablity and usurpability of their food resources. Recent evidence suggests that food may not be limiting for folivorous primates, and that male sexual coercion may be a more important influence on folivore social organization. To address the question, we collected focal data on the feeding behavior of adult females and males over 11 mo (September 2000-August 2001) on 2 groups: WW (n = 31–33 individuals) and B (n = 8–16 individuals). We also conducted phenological monitoring and a tree survey of the two-group home ranges to establish food availability and distribution. We used 2 behavioral or organism-defined indicators of feeding behavior to assess potential resource contestability: food site residence time and distance moved between food sites. The colobus fed on a high diversity of species, most of their food trees were not clumped in distribution, within-tree interfood distances were short, and food trees were large. The only condition associated with the potential for monopolization was low food tree density. However, low food tree density may be offset by the colobus’ use of large trees. Taken together, the ecological and behavioral indicators suggest the food resources of Colobus vellerosus had a low potential for monopolization. Our results also indicate mature leaves had the longest food site residence time, which may suggest they should be the most usurpable plant part, though their presumed low quality and high abundance probably counteracted the effect. The pattern implied the potential for direct feeding competition among Colobus vellerosus at Boabeng-Fiema was low and agonistic interactions over food are not expected. Instead, a group size effect on feeding efficiency should be a more predominant influence on feeding efficiency, if food is limiting for the species.     

Effect of Colony, Patch Distance, And Trajectory Sense on Movement Complexity in Foraging Ants

Animals display foraging trajectories when exploiting food patches. These displacements have been studied, particularly in organisms that forage from a central place. The complexity of a movement path may be analyzed by means of the fractal dimension, an index that estimates the tortuosity of a trajectory. In field experiments we studied the effect of trajectory sense, distance to the resource patch and colony on the movement complexity in a common ant of central Chile. We found that these factors and their interactions significantly affected the complexity of movement paths. We discuss whether mortality risk determines more linear trajectories when the resource patch is distant from the nest, and whether the information acquired from the environment by workers determines less complex return trips.