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How the local density of territorial animals responds to changes in food abundance will depend on the flexibility of territory size. Quantitative estimates of territory size over a broad range of food abundance are relatively rare because of the difficulty of measuring food abundance in the wild., Stream salmonids are an ideal model system for investigating flexibility in territory size, because food abundance can be quantified in the field and manipulated in the laboratory. We conducted a meta-analysis to test whether territory size decreases with increasing food abundance, and a mixed model analysis to test among three competing predictions: with increasing food abundance, territory size will be (1) fixed-the slope of a regression of log territory size vs. log food abundance = 0; (2) flexible and decreasing, as if individuals are defending a fixed amount of food-a slope = −1; and (3) initially compressible, but with an asymptotic minimum size-a slope between 0 and −1., We collected data from 16 studies that manipulated or measured food abundance while monitoring changes in territory size of young-of-the-year salmonids; 10 were experimental laboratory studies, whereas six were observational field studies., Overall, territory size decreased significantly with increasing food abundance; the weighted average correlation coefficient was −0.31. However, the estimated slope of the relationship between log territory size and log food abundance was only −0.23, significantly different from 0, and also significantly shallower than −1., Our estimated slope suggests that attempts to increase the density of territorial salmonids by increasing food abundance and reducing territory size will be inefficient; a 20-fold increase in food abundance would be required to double population density. Our analysis may also have implications for other species with a territorial mosaic social system-i.e. contiguous territories. In these social systems, social inertia will dampen any effects of changes in food abundance on the local density of settlers, compared to non-territorial species or those with non-contiguous territories. [ABSTRACT FROM AUTHOR]

There are several alternative hypotheses about the effects of territoriality, kinship and prey availability on individual carnivore distributions within populations. The first is the land-tenure hypothesis, which predicts that carnivores regulate their density through territoriality and temporal avoidance. The second is the kinship hypothesis, which predicts related individuals will be clumped within populations, and the third is the resource dispersion hypothesis, which suggests that resource richness may explain variable sociality, spatial overlap or temporary aggregations of conspecifics., Research on the socio-spatial organization of animals is essential in understanding territoriality, intra- and interspecific competition, and contact rates that influence diverse ecology, including disease transmission between conspecifics and courtship behaviours., We explored these hypotheses with data collected on a solitary carnivore, the cougar ( Puma concolor), from 2005 to 2012 in the Southern Yellowstone Ecosystem, Wyoming, USA. We employed 27 annual home ranges for 13 cougars to test whether home range overlap was better explained by land tenure, kinship, resource dispersion or some combination of the three., We found support for both the land tenure and resource dispersion hypotheses, but not for kinship. Cougar sex was the primary driver explaining variation in home range overlap. Males overlapped significantly with females, whereas the remaining dyads (F-F, M-M) overlapped significantly less. In support for the resource dispersion hypothesis, hunting opportunity (the probability of a cougar killing prey in a given location) was often higher in overlapping than in non-overlapping portions of cougar home ranges. In particular, winter hunt opportunity rather than summer hunt opportunity was higher in overlapping portions of female-female and male-female home ranges., Our results may indicate that solitary carnivores are more tolerant of sharing key resources with unrelated conspecifics than previously believed, or at least during periods of high resource availability. Further, our results suggest that the resource dispersion hypothesis, which is typically applied to social species, is applicable in describing the spatial organization of solitary carnivores. [ABSTRACT FROM AUTHOR]

Multiple approaches exist to model patterns of space use across species, among them resource selection analysis, statistical home-range modelling and mechanistic movement modelling. Mechanistic home-range models combine the benefits of these approaches, describing emergent territorial patterns based on fine-scale individual- or group-movement rules and incorporating interactions with neighbours and the environment. These models have not, to date, been extended to dynamic contexts., Using mechanistic home-range models, we explore meerkat ( Suricata suricatta) territorial patterns, considering scent marking, direct group interactions and habitat selection. We also extend the models to accommodate dynamic aspects of meerkat territoriality (territory development and territory shift)., We fit models, representing multiple working hypotheses, to data from a long-term meerkat study in South Africa, and we compare models using Akaike's and Bayesian Information Criteria., Our results identify important features of meerkat territorial patterns. Notably, larger groups do not seem to control larger territories, and groups apparently prefer dune edges along a dry river bed., Our model extensions capture instances in which 1) a newly formed group interacts more strongly with its parent groups over time and 2) a group moves its territory core out of aversive habitat. This extends our mechanistic modelling framework in previously unexplored directions. [ABSTRACT FROM AUTHOR]

Most animals experience temperature variations as they move through the environment. For ectotherms, in particular, temperature has a strong influence on habitat choice. While well studied at the species level, less is known about factors affecting the preferred temperature of individuals; especially lacking is information on how physiological traits are linked to thermal preference and whether such relationships are affected by factors such feeding history and growth trajectory., This study examined these issues in the common minnow Phoxinus phoxinus, to determine the extent to which feeding history, standard metabolic rate ( SMR) and aerobic scope ( AS), interact to affect temperature preference., Individuals were either: 1) food deprived (FD) for 21 days, then fed ad libitum for the next 74 days; or 2) fed ad libitum throughout the entire period. All animals were then allowed to select preferred temperatures using a shuttle-box, and then measured for SMR and AS at 10 °C, estimated by rates of oxygen uptake. Activity within the shuttle-box under a constant temperature regime was also measured., In both FD and control fish, SMR was negatively correlated with preferred temperature. The SMR of the FD fish was increased compared with the controls, probably due to the effects of compensatory growth, and so these growth-compensated fish preferred temperatures that were on average 2·85 °C cooler than controls fed a maintenance ration throughout the study. Fish experiencing compensatory growth also displayed a large reduction in activity. In growth-compensated fish and controls, activity measured at 10 °C was positively correlated with preferred temperature., Individual fish prefer temperatures that vary predictably with SMR and activity level, which are both plastic in response to feeding history and growth trajectories. Cooler temperatures probably allow individuals to reduce maintenance costs and divert more energy towards growth. A reduction in SMR at cooler temperatures, coupled with a decrease in spontaneous activity, could also allow individuals to increase surplus AS for coping with environmental stressors. In warming climates, however, aquatic ectotherms could experience frequent fluctuations in food supply with long-lasting effects on metabolic rate due to compensatory growth, while simultaneously having limited access to preferred cooler habitats. [ABSTRACT FROM AUTHOR]

Food availability is considered a major factor determining spacing behaviour in territorial species, especially for females. Theoretically, spatial overlap (considered the opposite of territoriality) and food availability are related in a nonlinear manner (hypothesized inverted- U function), with high overlap levels at the extremes of a food availability gradient and low overlap at intermediate levels of this gradient. Similar patterns are expected for encounter frequencies owing to its expected correlation with spatial overlap. However, these predictions have rarely been tested in highly structured social systems on a broad gradient of food availability, which implicitly requires experimental manipulation., We test these predictions in a solitary, territorial and trophic specialist, the Iberian lynx Lynx pardinus, taking advantage of a three-decade data set of spatial behaviour in different scenarios of food availability (i.e. rabbit density)., In contrast with expectations, home range overlap among resident females was low (median overlap index = 0·08, range 0-0·57) and core area overlap was nearly nil (median overlap index = 0, range 0-0·22) throughout the entire gradient of prey availability. Furthermore, spatial associations between pairs of females were negligible regardless marked variation in prey availability. Therefore, we did not find support for a model of flexible lynx territoriality driven by food availability., Our results suggest that the exclusive use of space in the Iberian lynx was not related to food. Lack of influence of prey availability on lynx territoriality may be adaptive to cope with the consequences of frequent drought-induced periods of prey scarcity or other disturbance typically affecting wild rabbit populations in Mediterranean environments. Thus, lynx would adopt an obstinate strategy of territoriality that consists in defending exclusive areas across a broad range of resource availability ensuring an exclusive access to the minimum amount of prey necessary for survival and eventually reproduction even during periods of prey scarcity. However, we found signs that territoriality was influenced by lynx density in a nonlinear fashion. Our results suggest the occurrence of population regulation through territoriality in this species. [ABSTRACT FROM AUTHOR]

Ants are ubiquitous ecosystem engineers and generalist predators and are able to affect ecological communities via both pathways. They are likely to influence any other terrestrial arthropod group either directly or indirectly caused by their high abundance and territoriality. We studied the impact of two ant species common in Central Europe, Myrmica rubra and Lasius niger, on an arthropod community. Colony presence and density of these two ant species were manipulated in a field experiment from the start of ant activity in spring to late summer. The experiment revealed a positive influence of the presence of one ant colony on densities of decomposers, herbivores and parasitoids. However, in the case of herbivores and parasitoids, this effect was reversed in the presence of two colonies. Generally, effects of the two ant species were similar with the exception of their effect on Braconidae parasitoid densities that responded positively to one colony of M. rubra but not of L. niger. Spider density was not affected by ant colony manipulation, but species richness of spiders responded positively to ant presence. This effect was independent of ant colony density, but where two colonies were present, spider richness was significantly greater in plots with two M. rubra colonies than in plots with one colony of each ant species. To test whether the positive ecosystem engineering effects were purely caused by modified properties of the soil, we added in an additional experiment (i) the soil from ant nests (without ants) or (ii) unmodified soil or (iii) ant nests (including ants) to experimental plots. Ant nest soil on its own did not have a significant impact on densities of decomposers, herbivores or predators, which were significantly, and positively, affected by the addition of an intact nest. The results suggest an important role of both ant species in the grassland food web, strongly affecting the densities of decomposers, herbivores and higher trophic levels. We discuss how the relative impact via bottom-up and top-down effects of ants depends on nest density, with a relatively greater top-down predatory impact at higher densities. [ABSTRACT FROM AUTHOR]

1. Population cycles are mostly thought to arise through extrinsic rather than intrinsic processes. However, in red grouse ( Lagopus lagopus scoticus), intrinsic male territoriality has been proposed as a driver of the cycles, possibly in conjunction with an extrinsic interaction with specialist parasitic worms. Here we examine how harvesting and environmental noise may also interact with territoriality to determine how grouse populations cycle. 2. A stochastic model of grouse dynamics based on the territoriality hypothesis is developed, including harvesting and the effects of nonterritorial birds on aggressiveness. Cycles are detected in 97% of populations simulated over realistic parameter ranges, and these exhibit similar statistical properties to those reported in studies of multiple grouse populations. As observed, cycle periods are shorter at higher breeding productivities. 3. The model demonstrates the destabilizing influence of delayed density-dependent territorial aggressiveness. Cycle amplitudes are higher when annual changes in aggression are larger and when nonterritorial males provoke greater aggression. Intriguingly, the model suggests how an interaction between aggressiveness and parasites may operate. It is known that males with high worm burdens show dramatic decreases in aggressiveness in the year following a peak in territoriality. When this is included in the model, via larger crashes in aggression, amplitudes are higher, despite a reduction in overall aggressiveness. 4. Environmental stochasticity interacts with territoriality to determine the form of the cycles, but this is mediated through its ‘colour’ or temporal autocorrelation. For example, uncorrelated white noise increases amplitudes, while autocorrelated red noise has the opposite effect. However, noise increases cycle periods whatever the colour. 5. Harvesting occurs before territorial competition. This reduces the pool of males competing for territories and so increases recruitment and population densities. However, crashes can then be more extreme so cycle amplitudes are higher. With harvesting at ~150% of current typical levels, which is within observed variation, the dynamics exhibit a sharp transition to a state where cyclicity is reduced, periods are shorter and amplitudes lower. 6. The model suggests that to understand regional variation in red grouse cycles, interactions between territoriality, productivity, harvesting and noise must be considered. [ABSTRACT FROM AUTHOR]

1. Empirical studies show that average growth of stream-dwelling salmon and trout often declines with increasing density in a characteristic concave relationship. However, the mechanisms that generate negative density–growth relationships in populations in natural streams are not certain. 2. In a recent study, Imre, Grant & Cunjak (2005; Journal of Animal Ecology, 74, 508–516) argue that density-dependent growth due to exploitative competition for prey causes the negative density–growth relationships for stream salmonids. They argue that the concave shape of empirical density–growth relationships is consistent with a simple model of exploitative competition and not consistent with interference competition for space. 3. We use a simple model to show that competition for space can yield concave density–growth relationships consistent with the empirical pattern when individuals compete for foraging sites that vary spatially in quality and lower-quality sites predominate. Thus, the predictions of the exploitative competition and spatial competition models overlap. 4. The shape of the density–growth relationship does not differentiate between candidate mechanisms underlying density-dependent growth for stream salmonids. Our results highlight the general problem with determining the mechanism driving an ecological process from patterns in observational data within the context of linking population demographics to habitat structure and animal behaviour. [ABSTRACT FROM AUTHOR]

1. Models on territory acquisition and tenure predict that territorial animals benefit by adjusting territorial defence behaviour to previous challenges they had experienced within the socially complex environment of communication networks. 2. Here, we addressed such issues of social cognition by investigating persisting effects of vocal contests on territory defence behaviour in nightingales Luscinia megarhynchos (Brehm). 3. Using interactive playback during nocturnal song of subjects, a rival was simulated to countersing either aggressively (by song overlapping) or moderately (by song alternating) from outside the subjects’ territory. Thereby, the time-specific singing strategy provided an experimentally controlled source of information on the motivation of an unfamiliar rival. 4. Expecting that nightingales integrate information with time, the same rival was simulated to return as a moderately singing intruder on the following morning. 5. The results show that the vigour with which male nightingales responded to the simulated intrusion of an opponent during the day depended on the nature of the nocturnal vocal interaction experienced several hours before. 6. Males that had received the song overlapping playback the preceding night approached the simulated intruder more quickly and closer and sang more songs near the loudspeaker than did males that had received a song alternating playback. 7. This adjustment of territory defence strategies depending on information from prior signalling experience suggests that integrating information with time plays an important part in territory defence by affecting a male's decision making in a communication network. [ABSTRACT FROM AUTHOR]

1. Costs of coexistence for species with indirect resource competition usually increase monotonically with competitor numbers. Very little is known though about the shape of the cost function for species with direct interference competition. 2. Here we report the results of an experiment with two vole species in artificial coexistence in large enclosures, where density of the dominant competitor species ( Microtus agrestis) was manipulated. Experimental populations of the subordinate vole species ( Clethrionomys glareolus) were composed of same aged individuals to study distribution of costs of coexistence with a dominant species within an age-cohort. 3. Survival and space use decreased gradually with increasing field vole numbers. Thus, responses to interference competition in our system appeared to be similar as expected from resource competition. The total number of breeders was stable. Reproductive characteristics such as the timing of breeding, and the litter size were not affected. In the single species enclosures a proportion of surviving individuals were not able to establish a breeding territory against stronger conspecifics. Under competition with heterospecifics such nonbreeders suffered high mortality, whereas the breeders survived. 4. Combined interference of dominant conspecifics and heterospecifics probably increased the frequency of aggressive interactions, social stress and mortality for the weaker individuals within a homogeneous age cohort of the subordinate competitor population. 5. Our results suggest, that in open systems where bank voles are outcompeted over the breeding season by faster reproducing field voles, animals able to establish a territory may be able to withstand competitor pressure, while nonbreeding bank vole individuals are forced to emigrate to suboptimal forest habitats. [ABSTRACT FROM AUTHOR]

1. In migratory birds males tend to arrive first on breeding grounds, except in sex-role reversed species. The two most common explanations are the rank advantage hypothesis, in which male–male competition for breeding sites drives stronger selection for early arrival in males than females, and the mate opportunity hypothesis, which relies on sexual selection, as early arrival improves prospects of mate acquisition more for males than for females. 2. To date, theoretical work has focused on selection for early arrival within a single sex, usually male. However, if fitness depends on territory quality, selection for early arrival should operate on both sexes. Here we use two independent modelling approaches to explore the evolution of protandry (male-first arrival) and protogyny (female-first arrival) under the rank advantage and mate opportunity hypotheses. 3. The rank advantage hypothesis, when operating alone, fails to produce consistent patterns of protandry, despite our assumption that males must occupy territories before females. This is because an individual of either sex benefits if it out-competes same-sex competitors. Rather than promoting protandry, the rank advantage mechanism can sometimes result in protogyny. Female–female competition is stronger than male–male competition early in the season, if females compete for a resource (territories occupied by males) that is initially less common than the resource of interest to males (unoccupied territories). 4. Our results support the mate opportunity hypothesis as an explanation of why protandry is the norm in migratory systems. Male-biased adult sex ratios and high levels of sperm competition (modelled as extra-pair young: EPY) both produce protandry as a result of sexual selection. Protogyny is only observed in our models with female-biased sex ratios and low EPY production. 5. We also show that the effects of sex ratio biases are much stronger than those of EPY production, explore the evidence for sex ratio biases and extra-pair paternity in migratory species and suggest future research directions. [ABSTRACT FROM AUTHOR]

1. Outbreaks of feral house mice, Mus domesticus, in Australia represent a fundamental failure of the behavioural control mechanisms of population density, as proposed in the hypothesis of self-regulation. 2. Mice have the potential to keep numbers in check via a suite of spacing behaviours; however, the self-regulation hypothesis implies that some social change occurs that permits the population to erupt. It also suggests that at different phases of an outbreak, distinct patterns of social activity are evident. 3. We compare predictions from two models encapsulating the self-regulation hypothesis as applied to feral house mice in south-eastern Australia. Each model may be distinguished by the timing of aggressiveness between mice that leads to a closed social system. We compare individual turnover, residency and territoriality in each sex and age cohort during the increase, peak and low phases of a population outbreak that peaked in 2001. 4. The activity of 438 mice was monitored via intensive mark–recapture trapping and an automated event recording system that detected the activity of 300 marked individuals at burrow entrances. 5. Our findings support the second model, which suggests that mice switch from an almost asocial structure at low densities to a territorial system as abundance increases. Adult females appear more likely than males or juveniles to make the significant social shift. The trigger for this change remains unclear and several alternative mechanisms are proposed. [ABSTRACT FROM AUTHOR]

1. The ecological factors that promote delayed dispersal of offspring in cooperatively breeding bird species are poorly understood. While single population studies have supported the view that natal dispersal is delayed as a consequence of lack of suitable breeding vacancies (ecological constraints hypothesis), recent theoretical models claim that habitat saturation cannot be the main factor leading to kin sociality. 2. The carrion crow ( Corvus corone ssp.) is an ideal model to investigate the ecology of delayed dispersal. The occurrence of kin sociality is highly variable among European populations, and it is known to be determined by environmental rather than genetic factors. Here we compare juvenile dispersal, territory turnover, habitat saturation (number of competitors per breeding vacancy), variability of territory quality and territoriality of two crow populations that differ in social organization (extensive kin sociality in Spain vs. social monogamy in Italy). 3. We found that philopatry occurred in the less competitive and less variable environment. A review of the information available in literature on other crow populations qualitatively supports this result. 4. In crows, juvenile philopatry seemed to be associated with breeders’ year-round residency in territories. This association is widespread among birds and it has been previously explained through an ecological constraints perspective, with residency slowing territory turnover and therefore augmenting saturation. Our data do not support this explanation. 5. We suggest that year-round residency might play a direct role in determining kin sociality in crows. In Spain, adults occupy and defend the territory year-round, providing the offspring with a place where they enjoy a preferential access to resources that may represent an incentive to stay. Conversely in Italy, where adults often abandon their territories after breeding, the natal site does not have any special value for the offspring that hence disperse. [ABSTRACT FROM AUTHOR]

Summary 1. Territory quality may affect individual fitness and contribute to density-dependent reproduction, with repercussions on population regulation. We investigated the probable causes and population consequences of spatio-temporal variations in territory quality, measured by occupancy, in eight black kite Milvus migrans Boddaert populations, one of them studied for 10 years (Lake Lugano) and the rest for 4–5 years. 2. Over a period of years, the occupation rate of territories varied from a random pattern. Some territories were preferred while others were avoided. On return from migration, males and females settled earlier on high-occupancy territories. 3. The positive association between territory occupancy and breeding performance held in all years of study at Lake Lugano, and in six of seven tested populations. As a result, high-occupancy territories contributed most of the young produced by each population. 4. The occupation rate of the overall 225 territories was related positively to food availability and negatively to mortality risk, measured as proximity to the nearest eagle owl Bubo bubo Linnaeus nest. 5. At the population level, spatial variation in mean occupancy was positively correlated with spatial variation in mean productivity, suggesting that mean occupancy could be used as a measure of overall habitat quality and population performance. 6. In the Lake Lugano area, a higher proportion of low quality territories was occupied in years of higher density and annual productivity was related negatively to its coefficient of variation. However, annual productivity was not related significantly to the proportion of low quality territories occupied, so support for the theory of site-dependent population regulation was only partial. 7. In a review of 22 studies of territory occupancy in 17 species, occupancy always deviated from a random pattern in species in which it was tested and was always correlated with productivity and/or with some other measure of territory quality. Our results confirm the importance of prioritizing conservation of high quality territories. Occupancy may be a reliable method of quality assessment, especially for populations in which not all territories are always occupied, or for species in which checking occupancy is easier than finding nests. [ABSTRACT FROM AUTHOR]

Summary 1. Based on kin selection theory, it has been predicted that close relatives should aggregate to accrue inclusive fitness advantages. However, studies of genetics have failed to find convincing evidence of kin association in wild populations of territorial fish such as juvenile salmonids, despite laboratory evidence of reduced aggression among siblings. In these genetic studies, calculations of the degree of kin association have been based on the assumption that the geographical distribution of individual fish is fixed over space and time. By comparison, recent work suggests that patterns of spatial distribution are fluid and that home ranges overlap considerably. Therefore, it is possible that benefits of kin selection depend not on the juxtaposition of fish per se , but on how they interact on the occasions when they associate. 2. To explore this issue, an experiment was conducted to test whether dominant fish in pairs of Atlantic salmon Salmo salar (L.) juveniles were more likely to share food in high quality feeding patches with kin than non-kin and also whether kin-biased intimidation from dominant to subordinate fish was sustained in the absence of the dominant individual. 3. Siblings shared high-quality feeding territories for twice as long as pairs of non-kin. The food intake of dominant fish did not vary with relatedness of subordinate conspecific. However, once dominant fish were satiated, subordinate individuals consumed a greater proportion of food items when paired with kin than non-kin. 4. When dominant fish were removed from half the pairs of salmon, subordinate fish that had been paired previously with dominant kin increased their feeding activity to greater levels than subordinate fish that had been paired previously with dominant non-kin. 5. These findings indicate that subordinate fish in wild populations may gain twofold foraging advantages by close association with dominant kin: (i) by increased territory sharing, and (ii) by reduced... [ABSTRACT FROM AUTHOR]

Summary 1. Ideal despotic distribution theory predicts that the quality of habitat controlled by territorial animals should vary depending on their competitive ability and the availability of resources. 2. In environments where resources have a patchy distribution, males that monopolize high quality territories may require less territory area than males in low quality areas. This has been a difficult premise to test in the wild owing to logistical constraints regarding manipulation of relevant resources and accurate measures of territory distributions. 3. We present results from an experimental test of ideal despotic distribution theory in a wild population of side-blotched lizards, Uta stansburiana (Baird and Girard). 4. We manipulated thermal resources on territories by shuttling rocks between dyads of neighbouring male territories. Manipulations created high quality territories by significantly increasing the variance in temperatures available for thermoregulation. 5. Experimentally improved quality territories (rock addition) became smaller after treatment, while reduced quality territories (rock removal) became larger. 6. Males on improved and reduced quality territories had equal numbers of females, resulting in higher densities of females on the smaller high quality territories. 7. Densities of the snake Masticophis flagellum , the dominant predator of Uta stansburiana , were higher on reduced quality territories. 8. Progeny released to experimental plots had significantly higher growth-rates and survival on experimentally improved sites relative to their neighbours on low quality territories. 9. Our results demonstrate both the ecological factors that drive the ideal despotic distribution, and the fitness consequences of high and low quality territories to lizards. [ABSTRACT FROM AUTHOR]