Spatial pattern changes of grazing cattle population I

A cattle population presents three spatial patterns according to their behaviour. These patterns are as follows: the random spatial pattern which occurs when the cattle move during grazing (feeding), the aggregated spatial pattern which occurs when they rest, and the regular or quasi-regular spatial pattern which occurs when they separate into two or more subgroups. In this study, these three states of behaviour were represented by a mathematical model, and a method to test the pattern statistically was derived.     

Modelling the Spatial Behaviour of a Tropical Tuna Purse Seine Fleet

Industrial tuna fisheries operate in the Indian, Atlantic and Pacific Oceans, but concerns over sustainability and environmental impacts of these fisheries have resulted in increased scrutiny of how they are managed. An important but often overlooked factor in the success or failure of tuna fisheries management is the behaviour of fishers and fishing fleets. Uncertainty in how a fishing fleet will respond to management or other influences can be reduced by anticipating fleet behaviour, although to date there has been little research directed at understanding and anticipating the human dimension of tuna fisheries. The aim of this study was to address gaps in knowledge of the behaviour of tuna fleets, using the Indian Ocean tropical tuna purse seine fishery as a case study. We use statistical modelling to examine the factors that influence the spatial behaviour of the purse seine fleet at broad spatiotemporal scales. This analysis reveals very high consistency between years in the use of seasonal fishing grounds by the fleet, as well as a forcing influence of biophysical ocean conditions on the distribution of fishing effort. These findings suggest strong inertia in the spatial behaviour of the fleet, which has important implications for predicting the response of the fleet to natural events or management measures (e.g., spatial closures).     

Seasonal changes in hippocampus size and spatial behaviour in mammals and birds

Hippocampus is involved in processing of environmental spatial information, and its size is known to correlate positively with spatial abilities in mammals and birds. Comparisons between species suggest that amount of spatial information processed (the mean area of home range in particular) is related with hippocampus size. Do seasonal and age changes in hippocampus size correlate with seasonal dynamics of spatial behaviour during ontogenesis? The data obtained through observational and experimental studies confirm the possibility that hippocampus size may be subjected to adaptive modifications along with cyclic changes in spatial behavior. In course of seasonal dynamics, strong positive correlation was found between hippocampus mass, home range size, and mobility of small mammals. Recently, first facts demonstrating seasonal changes of hippocampus and spatial behaviour (in connection with food-storing and brood parasitism) were found in birds. A lot of facts obtained for different taxonomical groups shows parallel seasonal changes in spatial behaviour and morphology of brain region functionally related to such behaviour. Thus, in adult birds and mammals, not only behaviour but also brain structure is phenotypically flexible in response to seasonally changing environment. Morphophysiological mechanisms of hippocampus seasonal changes are also discussed.     

Spatial-pattern analysis in a territorial spider: evidence for multi-scale effects

Territorial animals maintain a certain distance to neighbouring conspecifics, presumably leading to a regular spatial pattern through social spacing. Nevertheless, most animal populations are assumed to show aggregation at certain distance ranges, reflecting the scale dependency of spatial patterns. Leucorchestris arenicola (Araneae: Sparassidae) is a burrow-living spider species that shows territorial behaviour against conspecifics. A multi-scale approach in spatial analysis revealed that territory owners had fewer neighbours than expected under spatial randomness at distances up to 6 m. Behavioural field experiments showed that territory owners were able to perceive and react to burrow constructing neighbours up to at least 4 m distance from their own burrow. At larger distances individuals were often more aggregated than expected under spatial randomness. Analysing adult and immature relationships showed attraction between different development stages at small distances and avoidance at larger distances. The analysis reveals diverse spatial patterns in a territorial and cannibalistic species, showing that both behaviour and environment affect pattern development at different distances. The study outlines the importance of multi-scale approaches for spatial analysis and the need for accompanying experiments to facilitate the interpretation of results.     

Lateral position preference in grazing feral horses

Behavioural lateralisation is an effective way for animals to manage daily tasks by specialising behaviour to either side of the body. Many types of lateralisation are linked to the function of each brain hemisphere. Lateralisation of monitoring behaviour in mother–infant relationships occurs in a wide range of mammals, where infants frequently use their left eye to monitor their mother. However, few studies have focused on this type of spatial relationships among adults in daily life, such as during foraging. The present study focused on monitoring adult feral horse behaviour using quantitative analysis of spatial relationships, using drone technology. We found that horses form a localised spatial relationship with their nearest neighbour. Specifically, the nearest neighbour was located to the left rear of a target individual significantly more frequently than to the right rear. Furthermore, the nearest neighbour was less frequently located behind a target individual. We propose that this relationship is caused by a left-eye preference, because information via the left eye predominantly proceeds to the right hemisphere, which is dominant for social processing.     

Using ecology to guide the study of cognitive and neural mechanisms of different aspects of spatial memory in food-hoarding animals

Understanding the survival value of behaviour does not tell us how the mechanisms that control this behaviour work. Nevertheless, understanding survival value can guide the study of these mechanisms. In this paper, we apply this principle to understanding the cognitive mechanisms that support cache retrieval in scatter-hoarding animals. We believe it is too simplistic to predict that all scatter-hoarding animals will outperform non-hoarding animals on all tests of spatial memory. Instead, we argue that we should look at the detailed ecology and natural history of each species. This understanding of natural history then allows us to make predictions about which aspects of spatial memory should be better in which species. We use the natural hoarding behaviour of the three best-studied groups of scatter-hoarding animals to make predictions about three aspects of their spatial memory: duration, capacity and spatial resolution, and we test these predictions against the existing literature. Having laid out how ecology and natural history can be used to predict detailed cognitive abilities, we then suggest using this approach to guide the study of the neural basis of these abilities. We believe that this complementary approach will reveal aspects of memory processing that would otherwise be difficult to discover.     

State-space models of individual animal movement

Detailed observation of the movement of individual animals offers the potential to understand spatial population processes as the ultimate consequence of individual behaviour, physiological constraints and fine-scale environmental influences. However, movement data from individuals are intrinsically stochastic and often subject to severe observation error. Linking such complex data to dynamical models of movement is a major challenge for animal ecology. Here, we review a statistical approach, state-space modelling, which involves changing how we analyse movement data and draw inferences about the behaviours that shape it. The statistical robustness and predictive ability of state-space models make them the most promising avenue towards a new type of movement ecology that fuses insights from the study of animal behaviour, biogeography and spatial population dynamics.     

Self-organization, scale and stability in a spatial predator-prey interaction

Simple predator-prey models often predict extreme instability in interactions where the prey are depressed well below their carrying capacity. Although the behaviour of some laboratory systems conforms to this pattern, field and mesocosm studies generally show prolonged co-existence of prey and predator. Prominent among the possible causes of this discrepancy are the effects of spatial heterogeneity. In this paper we show that both discrete and continuous representations of the spatial Rosenzweig-McArthur model with immobile prey can be stabilized by self-organized prey heterogeneity. This concordance of behaviour closely parallels that which we have previously established in the context of invasion waves. We use the continuous model variant to calculate the characteristic spatial scales of the self-organized structures. The discrete variant forms the basis of a simulation study demonstrating the variety of stable structures and elucidating their relation to the history of the system. We note that all stable prey distributions take the form of a network of occupied patches separated by prey-free regions, and liken the process which generates such assemblages to the formation of a landscape mozaic.     

A transversely isotropic hyperelastic constitutive model of the PDL. Analytical and computational aspects

This study describes the development of a constitutive law for the modelling of the periodontal ligament (PDL) and its practical implementation into a commercial finite element code. The constitutive equations encompass the essential mechanical features of this biological soft tissue: non-linear behaviour, large deformations, anisotropy, distinct behaviour in tension and compression and the fibrous characteristics. The approach is based on the theory of continuum fibre-reinforced composites at finite strain where a compressible transversely isotropic hyperelastic strain energy function is defined. This strain energy density function is further split into volumetric and deviatoric contributions separating the bulk and shear responses of the material. Explicit expressions of the stress tensors in the material and spatial configurations are first established followed by original expressions of the elasticity tensors in the material and spatial configurations. As a simple application of the constitutive model, two finite element analyses simulating the mechanical behaviour of the PDL are performed. The results highlight the significance of integrating the fibrous architecture of the PDL as this feature is shown to be responsible for the complex strain distribution observed.     

A Transversely Isotropic Hyperelastic Constitutive Model of the PDL. Analytical and Computational Aspects

This study describes the development of a constitutive law for the modelling of the periodontal ligament (PDL) and its practical implementation into a commercial finite element code. The constitutive equations encompass the essential mechanical features of this biological soft tissue: non-linear behaviour, large deformations, anisotropy, distinct behaviour in tension and compression and the fibrous characteristics. The approach is based on the theory of continuum fibre-reinforced composites at finite strain where a compressible transversely isotropic hyperelastic strain energy function is defined. This strain energy density function is further split into volumetric and deviatoric contributions separating the bulk and shear responses of the material. Explicit expressions of the stress tensors in the material and spatial configurations are first established followed by original expressions of the elasticity tensors in the material and spatial configurations. As a simple application of the constitutive model, two finite element analyses simulating the mechanical behaviour of the PDL are performed. The results highlight the significance of integrating the fibrous architecture of the PDL as this feature is shown to be responsible for the complex strain distribution observed.     

Diel variation in feeding and vertical distribution of ten co-occurring fish species: consequences for resource partitioning

The diel variations in feeding behaviour and vertical distribution were determined for ten species of a tropical fish community in a shallow SE Sri Lankan reservoir. The fish community consisted of two introduced exotic tilapias and eight indigenous riverine species including five cyprinids, one clarid, one hemiramphid and one gobid. Multi-mesh gillnets dividing the water column in six depth strata were set with a six-hour interval throughout four 24-hour periods. The study of the diel variation in feeding behaviour was based on analysis of catch data, gut contents and gut fullness per setting time. The diel variation in vertical distribution was determined per species for each of the six depth-strata. Feeding behaviour and vertical distribution differed significantly among species and for most species throughout the 24-hour period. Feeding behaviour and vertical distribution were correlated. The status of whether or not a fish is feeding determines if the trophic and spatial resource dimensions are interrelated. Resource partitioning along the spatial dimension may reduce exploitative competition at the time of feeding, whereas it may limit interference competition or predation during non-feeding periods. This revised version was published online in July 2006 with corrections to the Cover Date.     

An individual-oriented model on the emergence of support in fights, its reciprocation and exchange

Complex social behaviour of primates has usually been attributed to the operation of complex cognition. Recently, models have shown that constraints imposed by the socio-spatial structuring of individuals in a group may result in an unexpectedly high number of patterns of complex social behaviour, resembling the dominance styles of egalitarian and despotic species of macaques and the differences between them. This includes affiliative patterns, such as reciprocation of grooming, grooming up the hierarchy, and reconciliation. In the present study, we show that the distribution of support in fights, which is the social behaviour that is potentially most sophisticated in terms of cognitive processes, may emerge in the same way. The model represents the spatial grouping of individuals and their social behaviour, such as their avoidance of risks during attacks, the self-reinforcing effects of winning and losing their fights, their tendency to join in fights of others that are close by (social facilitation), their tendency to groom when they are anxious, the reduction of their anxiety by grooming, and the increase of anxiety when involved in aggression. Further, we represent the difference in intensity of aggression apparent in egalitarian and despotic macaques. The model reproduces many aspects of support in fights, such as its different types, namely, conservative, bridging and revolutionary, patterns of choice of coalition partners attributed to triadic awareness, those of reciprocation of support and 'spiteful acts' and of exchange between support and grooming. This work is important because it suggests that behaviour that seems to result from sophisticated cognition may be a side-effect of spatial structure and dominance interactions and it shows that partial correlations fail to completely omit these effects of spatial structure. Further, the model is falsifiable, since it results in many patterns that can easily be tested in real primates by means of existing data.     

Predation risk and the evolutionary ecology of reproductive behaviour

A large literature exists on the effects of predation risk on foraging and survival-related behaviours. In contrast, with some notable exceptions, relatively few theoretical or experimental studies have examined the effects of predation risk on reproductive behaviours. Existing literature on risk and reproductive behaviour and suggestion directions for future study are reviewed. In particular: (1) effects on predation risk on mating behaviour; (2) the influence of spatial patchiness on interactions between risk and reproductive behaviour; (3) the potential influence of multi-species interactions on the effects of predation risk on mating dynamics; (4) the importance of looking at sets of inter-related antipredator traits; and (5) some effects of predation risk on prey population patterns due to changes in prey reproductive behaviour are discussed. To illustrate various points examples involving fish and other aquatic organisms are used.     

The automated analysis of clustering behaviour of piglets from thermal images in response to immune challenge by vaccination

An automated method of estimating the spatial distribution of piglets within a pen was used to assess huddling behaviour under normal conditions and during a febrile response to vaccination. The automated method was compared with a manual assessment of clustering activity. Huddling behaviour was partly related to environmental conditions and clock time such that more huddling occurred during the night and at lower ambient air temperatures. There were no positive relationships between maximum pig temperatures and environmental conditions, suggesting that the narrow range of air temperatures in this study was not a significant factor for pig temperature. Spatial distribution affected radiated pig temperature measurements by IR thermography. Higher temperatures were recorded in groups of animals displaying huddling behaviour. Huddling behaviour was affected by febrile responses to vaccination with increased huddling occurring 3 to 8 h post-vaccination. The automated method of assessing spatial distribution from an IR image successfully identified periods of huddling associated with a febrile response, and to changing environmental temperatures. Infrared imaging could be used to quantify temperature and behaviour from the same images.     

Use of space and habitat selection by roe deer Capreolus capreolus in a Mediterranean coastal area: how does woods landscape affect home range?

The home-range dynamics and habitat selection of nine roe deer were studied from March 1994 to August 1994 in the Maremma Natural Park along the Tyrrhenian coast of Italy. The habitat was highly fragmented, with open agricultural fields prevailing in the study area (57%); the climate was Mediterranean. Data on spatial behaviour were collected by radio-tracking techniques. Habitat selection and structure were investigated by compositional and landscape analysis, both within the study area and within the home ranges. Animals of our sample showed spatial-use patterns varying from stationary to roaming. Stationary individuals used small home ranges while roaming ones moved, especially during the reproductive period in July and August. The percentage and structure of woodlands influenced the size of home ranges and the behaviour of males: stationary males used large amounts of woodlands within their home ranges and showed a territorial behaviour whereas males that used a high percentage of fields showed wider home ranges even during the territorial period. Females seemed to be less influenced by the presence and patch-structure of woodland within their home range. Landscape structure and habitat composition seemed to be important factors influencing the spatial behaviour of this roe deer population.     

European starlings recognize the location of robotic conspecific attention

Looking where others are allocating attention can facilitate social interactions by providing information about objects or locations of interest. We asked whether European starlings follow the orientation behaviour of conspecifics owing to their highly gregarious behaviour. Starlings reoriented their attention to follow that of a robot around a barrier more often than when the robot''s attention was directed elsewhere. This is the first empirical evidence of reorienting in response to conspecific attention in a songbird. Starlings may use this behaviour to obtain fine-tuned spatial information from conspecifics (e.g. direction of predator approach, spatial location of food patches), enhancing group cohesion.     

Intraspecific variation in herbivore‐induced plant volatiles influences the spatial range of plant–parasitoid interactions

Chemical information influences the behaviour of many animals, thus affecting species interactions. Many animals forage for resources that are heterogeneously distributed in space and time, and have evolved foraging behaviour that utilizes information related to these resources. Herbivore‐induced plant volatiles (HIPVs), emitted by plants upon herbivore attack, provide information on herbivory to various animal species, including parasitoids. Little is known about the spatial scale at which plants attract parasitoids via HIPVs under field conditions and how intraspecific variation in HIPV emission affects this spatial scale. Here, we investigated the spatial scale of parasitoid attraction to two cabbage accessions that differ in relative preference of the parasitoid Cotesia glomerata when plants were damaged by Pieris brassicae caterpillars. Parasitoids were released in a field experiment with plants at distances of up to 60 m from the release site using intervals between plants of 10 or 20 m to assess parasitism rates over time and distance. Additionally, we observed host‐location behaviour of parasitoids in detail in a semi‐field tent experiment with plant spacing up to 8 m. Plant accession strongly affected successful host location in field set‐ups with 10 or 20 m intervals between plants. In the semi‐field set‐up, plant finding success by parasitoids decreased with increasing plant spacing, differed between plant accessions, and was higher for host‐infested plants than for uninfested plants. We demonstrate that parasitoids can be attracted to herbivore‐infested plants over large distances (10 m or 20 m) in the field, and that stronger plant attractiveness via HIPVs increases this distance (up to at least 20 m). Our study indicates that variation in plant traits can affect attraction distance, movement patterns of parasitoids, and ultimately spatial patterns of plant–insect interactions. It is therefore important to consider plant‐trait variation in HIPVs when studying animal foraging behaviour and multi‐trophic interactions in a spatial context.     

Dispersal Patterns,Active Behaviour,and Flow Environment during Early Life History of Coastal Cold Water Fishes

During the pelagic larval phase, fish dispersal may be influenced passively by surface currents or actively determined by swimming behaviour. In situ observations of larval swimming are few given the constraints of field sampling. Active behaviour is therefore often inferred from spatial patterns in the field, laboratory studies, or hydrodynamic theory, but rarely are these approaches considered in concert. Ichthyoplankton survey data collected during 2004 and 2006 from coastal Newfoundland show that changes in spatial heterogeneity for multiple species do not conform to predictions based on passive transport. We evaluated the interaction of individual larvae with their environment by calculating Reynolds number as a function of ontogeny. Typically, larvae hatch into a viscous environment in which swimming is inefficient, and later grow into more efficient intermediate and inertial swimming environments. Swimming is therefore closely related to length, not only because of swimming capacity but also in how larvae experience viscosity. Six of eight species sampled demonstrated consistent changes in spatial patchiness and concomitant increases in spatial heterogeneity as they transitioned into more favourable hydrodynamic swimming environments, suggesting an active behavioural element to dispersal. We propose the tandem assessment of spatial heterogeneity and hydrodynamic environment as a potential approach to understand and predict the onset of ecologically significant swimming behaviour of larval fishes in the field.     

Elucidating the significance of spatial memory on movement decisions by African savannah elephants using state–space models

Spatial memory facilitates resource acquisition where resources are patchy, but how it influences movement behaviour of wide-ranging species remains to be resolved. We examined African elephant spatial memory reflected in movement decisions regarding access to perennial waterholes. State–space models of movement data revealed a rapid, highly directional movement behaviour almost exclusively associated with visiting perennial water. Behavioural change point (BCP) analyses demonstrated that these goal-oriented movements were initiated on average 4.59 km, and up to 49.97 km, from the visited waterhole, with the closest waterhole accessed 90% of the time. Distances of decision points increased when switching to different waterholes, during the dry season, or for female groups relative to males, while selection of the closest waterhole decreased when switching. Overall, our analyses indicated detailed spatial knowledge over large scales, enabling elephants to minimize travel distance through highly directional movement when accessing water. We discuss the likely cognitive and socioecological mechanisms driving these spatially precise movements that are most consistent with our findings. By applying modern analytic techniques to high-resolution movement data, this study illustrates emerging approaches for studying how cognition structures animal movement behaviour in different ecological and social contexts.     

Spatial movement of adult leopard seals (Hydrurga leptonyx) in Prydz Bay, Eastern Antarctica

This is the first study that has used satellite telemetry to assess the spatial behaviour of adult leopard seals. Satellite tags on 11 leopard seals transmitted between 29 days and 282 days. Swim speeds, distances swum per day and distances from tagging site were significantly different among individuals and seasons. Swim speeds ranged from 0.004 km/h to 10.86 km/h; distances swum per day from 0 km/day to 150 km/day; and the maximum distances from tagging site ranged from 33.30 km to 319.97 km. Rather than moving north-south with the ice most seals remained within a 50 km radius of their tagging site from 11 days to 97 days. The relatively sedentary movement of the leopard seals was unexpected, particularly the movement of animals over winter, which although slightly offshore did not reflect the usual northward winter migration described for the leopard seal. But traditionally, the leopard seals spatial habits have been described from sightings of animals at higher latitudes. These are generally younger animals and their behaviour may not be representative of the adults. This study has focused on adult females and animals at the extreme southerly range of the leopard seal. This highlights the importance of understanding and reporting age structure and distribution when discussing animal spatial behaviour.