Due to its conspicuous manifestations and its capacity to shape the configuration and dynamics of wild populations, territorial behavior has long intrigued ecologists. Territoriality and other animal interactions in situ have traditionally been studied via direct observations and telemetry. Here, we explore whether noninvasive genetic sampling, which is increasingly supplementing traditional field methods in ecological research, can reveal territorial behavior in an elusive carnivore, the wolverine (Gulo gulo). Using the locations of genotyped wolverine scat samples collected annually over a period of 12 years in central Norway, we test three predictions: (1) male home ranges constructed from noninvasive genetic sampling data are larger than those of females, (2) individuals avoid areas used by other conspecifics of the same sex (intrasexual territoriality), and (3) avoidance of same-sex territories diminishes or disappears after the territory owner's death. Each of these predictions is substantiated by our results: sex-specific differences in home range size and intrasexual territoriality in wolverine are patently reflected in the spatial and temporal configuration of noninvasively collected genetic samples. Our study confirms that wildlife monitoring programs can utilize the spatial information in noninvasive genetic sampling data to detect and quantify home ranges and social organization.

In solitary carnivorous mammals, territoriality is assumed to benefit male fitness by ensuring the exclusivity of matings within territories via mate guarding and female defence. However, this hypothesis remains empirically untested. Here, we examined this hypothesis for solitary territorial carnivores using the Iberian lynx (Lynx pardinus) as a case study. We expected that territorial males sire all litters born within their territories, translating into the absence of multi-paternity cases within the same litter. We analysed parentage in 43 kittens, belonging to 20 different litters. For 42 kittens, a father could be assigned using microsatellites and always coincided with the individual holding the territory. For 16 kittens from 10 litters for which we also had information on SNPs, paternity assignments coincided with microsatellites, except for a litter (two kittens) from the same litter for which a different male was assigned, but the territorial male could not be excluded. Our results indicated that multi-paternity in the Iberian lynx must be a rare event, and that territorial males sire all litters born from the females with which they share territories. We propose that both the low number of mature individuals in the lynx population and the fact that female oestrus is induced by male presence may explain results.

Bacteria display social behavior and establish cooperative or competitive interactions in the niches they occupy. The human skin is a densely populated environment where many bacterial species live. Thus, bacterial inhabitants are expected to find a balance in these interactions, which eventually defines their spatial distribution and the composition of our skin microbiota. Unraveling the physiological basis of the interactions between bacterial species in organized environments requires reductionist analyses using functionally relevant species. Here, we study the interaction between two members of our skin microbiota, Bacillus subtilis and Staphylococcus epidermidis. We show that B. subtilis actively responds to the presence of S. epidermidis in its proximity by two strategies: antimicrobial production and development of a subpopulation with migratory response. The initial response of B. subtilis is production of chlorotetain, which degrades the S. epidermidis at the colony level. Next, a subpopulation of B. subtilis motile cells emerges. Remarkably this subpopulation slides towards the remaining S. epidermidis colony and engulfs it. A slow response back from S. epidermidis cells give origin to resistant cells that prevent both attacks from B. subtilis. We hypothesized that this niche conquering and back-down response from B. subtilis and S. epidermidis, respectively, which resembles other conflicts in nature as the ones observed in animals, may play a role in defining the presence of certain bacterial species in the specific microenvironments that these bacteria occupy on our skin.

After entry into lymph nodes (LNs), B cells migrate to follicles, whereas T cells remain in the paracortex, with each lymphocyte type showing apparently random migration within these distinct areas. Other than chemokines, the factors contributing to this spatial segregation and to the observed patterns of lymphocyte movement are poorly characterized. By combining confocal, electron, and intravital microscopy, we showed that the fibroblastic reticular cell network regulated naive T cell access to the paracortex and also supported and defined the limits of T cell movement within this domain, whereas a distinct follicular dendritic cell network similarly served as the substratum for movement of follicular B cells. These results highlight the central role of stromal microanatomy in orchestrating cell migration within the LN.

Road ecology has traditionally focused on the impact of in-situ and functional roads on wildlife. However, road construction also poses a major, yet understudied, threat and the implications for key aspects of animal behaviour are unknown. Badgers (Meles meles) have been implicated in the transmission of tuberculosis to cattle. There are concerns that environmental disturbances, including major road construction, can disrupt badger territoriality, promoting the spread of the disease to cattle. To address these knowledge gaps the ranging behaviour of a medium-density Irish badger population was monitored using GPS-tracking collars before, during, and after a major road realignment project that bisected the study area. We estimated badgers' home range sizes, nightly distances travelled, and the distance and frequency of extra-territorial excursions during each phase of the study and quantified any changes to these parameters. We show that road construction had a very limited effect on ranging behaviour. A small increase in nightly distance during road construction did not translate into an increase in home range size, nor an increase in the distance or frequency of extra-territorial excursions during road construction. In addition, suitable mitigation measures to prevent badger deaths appeared to ensure that normal patterns of ranging behaviour continued once the new road was in place. We recommend that continuous badger-proof fencing be placed along the entire length of new major roads, in combination with appropriately sited underpasses. Our analysis supports the view that road construction did not cause badgers to change their ranging behaviour in ways likely to increase the spread of tuberculosis.

Competition is considered to rely on the value attributed to resources by animals, but the influence of extrinsic stressors on this value remains unexplored. Although natural or anthropogenic environmental stress often drives decreased competition, assumptions that this relies on resource devaluation are without formal evidence. According to theory, physiological or perceptual effects may influence contest behaviour directly, but motivational changes due to resource value are expected to manifest as behavioural adjustments only in interaction with attainment costs and resource benefits. Thus, we hypothesise that stressor-induced resource devaluations will impose greater effects when attainment costs are high, but not when resource benefits are higher. Noise may elicit such effects because it impacts the acoustic environment and imposes physiological and behavioural costs to animals. Therefore, we manipulated the acoustic environment using playbacks of artificial noise to test our hypotheses in the territorial male Siamese fighting fish, Betta splendens.

Interest in control methods for invasive wild pigs (Sus scrofa) has increased due to their range expansion, population growth, and an improved understanding of their destructive ecological and economic effects. Recent technological advances in traps for control of pig populations facilitate capture of entire social groups (sounders), but the efficacy of "whole-sounder" trapping strategies is heavily dependent on the degree of territoriality among sounders, a topic little research has explored. We assessed territoriality in wild pig sounders on the Savannah River Site, South Carolina, USA, and examined whether availability of food resources provided by a municipal-waste landfill affected among-sounder territoriality. We estimated utilization distribution overlap and dynamic interactions among 18 neighboring sounders around a landfill. We found that although neighboring sounders overlapped in space, intensity of use in shared areas was uniformly low, indicating territorial behavior. Neighbors tended to share slightly more space when closer to the landfill waste cells, indicating availability of a super-abundant resource somewhat weakens the degree of territoriality among sounders. Nevertheless, we conclude that sounders behaved in a generally territorial manner, and we discuss implications for whole-sounder trapping programs, particularly near concentrated resources such as landfills and crop fields.

The connection between testosterone and territoriality in free-living songbirds has been well studied in a reproductive context, but less so outside the breeding season. To assess the effects of seasonal androgenic action on territorial behavior, we analyzed vocal and non-vocal territorial behavior in response to simulated territorial intrusions (STIs) during three life-cycle stages in free-living male black redstarts: breeding, molt and nonbreeding. Concurrently, we measured changes in circulating testosterone levels, as well as the mRNA expression of androgen and estrogen receptors and aromatase in the preoptic, hypothalamic and song control brain areas that are associated with social and vocal behaviors.

Novel ecosystems have emerged through human intervention and are rapidly expanding around the world. Whether they can support animal wildlife has generated considerable controversy. Here we developed a new approach to evaluate the ability of a novel forest ecosystem, dominated by the exotic tree species Eucalyptus globulus, to support animal wildlife in the medium and long term. To evaluate this ability, we took advantage of the fact that species territory size decreases with increasing habitat quality, and we used territoriality of a raptor guild composed of Northern Goshawk (Accipiter gentilis), Eurasian Sparrowhawk (A. nisus) and Common Buzzard (Buteo buteo) as indicator. We compared the territoriality of these species in the novel ecosystem with that in other ecosystems found in the literature. Average distances between con-specifics in the novel ecosystem were similar, or even shorter, than those in other ecosystems. Average distances between Goshawk con-specifics were among the shortest described in the literature. All three species nested preferably in mixed stands abundant in large exotic trees, with high structural complexity and abundance of native species within the stand. Key factors supporting this diverse and dense raptor community were the special forest management system implemented in the study area and the agricultural matrix located close to forest plantations that complements the supply of prey. Our results suggest that forest management that promotes a complex and suitable forest structure can increase the ability of novel forest ecosystems to support wildlife biodiversity, particularly a diverse nesting community of forest-dwelling raptors and their preys. The results further suggest the suitability of territoriality for assessing this potential of novel ecosystems.

Proteus mirabilis is a pathogenic Gram-negative bacterium characterized by its ability to swarm across surfaces, which frequently leads to colonization of the urinary tract and causes severe infections. P. mirabilis strains are also well known from their self-recognition phenomenon, referred to as Dienes phenomenon. In this study, we present novel aspect of self-recognition, which is a hierarchy in terms of strains territoriality. We report the draft genome sequences of P. mirabilis K1609 and K670 strains exhibiting the strongest and the weakest territoriality, respectively. Our results indicated that K1609 is closely related to strain BB2000, a model system for self-recognition, comparing with the K670. We annotated genes associated with recognition of kin and swarming initiation control and indicated polymorphisms by which observed differences in territoriality might results from. The phenotypic and genomic features of both strains reveal their application as a model organisms for studying not only the mechanisms of kin-recognition but also strains territoriality, thus providing new approach to the phenomenon. Availability of these genome sequences may facilitate understanding of the interactions between P. mirabilis strains.

While developmental consequences of parental investment on species-typical social behaviors has been extensively characterized in same-sex parent-offspring interactions, the impact of opposite-sex relationships is less clear. In the bi-parental California mouse (Peromyscus californicus), paternal retrieval behavior induces territorial aggression and the expression of arginine vasopressin (AVP) in adult male offspring. Although similar patterns of territorially emerge among females, the sexually dimorphic AVP system has not been considered since it is generally thought to regulate male-typical behavior. However, we recently demonstrated that male and female P. californicus offspring experience increases in plasma testosterone following paternal retrieval. Since AVP expression is androgen-dependent during development, we postulate that increases in AVP expression may accompany territoriality in female, as well as male offspring. To explore this aim, adult P. californicus offspring that received either high or low levels of paternal care (retrievals) during early development were tested for territoriality and immunohistochemical analysis of AVP within the bed nucleus of the stria terminalis (BNST), paraventricular nucleus (PVN), and supraoptic nucleus (SON). Consistent with previous studies, high care offspring were more aggressive than low care offspring. Moreover, high care offspring had significantly more AVP immunoreactive (AVP-ir) cells within the BNST than low care offspring. This pattern was observed within female as well as male offspring, suggesting an equally salient role for paternal care on female offspring physiology. Regardless of early social experience, sex differences in AVP persisted in the BNST, with males having greater expression than females.

Substrate vibration communication is displayed by a variety of insects that rely on silk for shelter. Such signaling is often associated with territoriality and social interactions. The goal in this study was to explore the use of substrate vibration by subsocial insects of the little-studied order Embioptera (also known as Embiidina). Antipaluria urichi (Saussure) (Embioptera: Clothodidae) from Trinidad and Tobago, a large embiopteran, exhibits maternal care and facultatively colonial behavior. Previous observations suggested that they were aggressive while guarding eggs but gregarious when not. Egg-guarders in particular have been observed shaking and lunging their bodies, but to date these putative signals have not been recorded nor were their contexts known. Staged interactions were conducted in the laboratory using residents that had established silk domiciles enveloping piezo-electric film used to detect vibrations. Predictions from two competing hypotheses, the maternal territoriality hypothesis and the group cohesion hypothesis, were erected to explain the occurrence of signaling. Experiments pitted pre-reproductive and egg-guarding residents against female and male intruders, representing social partners that ranged from potentially threatening to innocuous or even helpful. Behavioral acts were identified and scored along with associated substrate vibrations, which were measured for associated body movements, duration, and frequency spectra. Signals, sorted by the distinct actions used to generate them, were lunge, shake, push up, and snapback. Egg-guarding females produced most signals in response to female intruders, a result that supported the maternal territoriality hypothesis. Female intruders generally responded to such signaling by moving away from egg-guarding residents. In contrast, pre-reproductive residents did not signal much, and intruders settled beside them. Theme software was used to analyze the behavioral event recordings to seek patterns over time and their association with signals. Long patterns of behavioral acts were associated with shakes, lunges, and push-ups, indicating that interactions were occurring between the residents and intruders as would be expected when communication occurs. The value of Theme software, as well as the relationship between signaling by A. urichi and the risks and benefits of coloniality, are discussed.

Territoriality is an adaptive behavioral trait that is important for animal's fitness and there still remains much to learn about the proximate mechanisms underlying the development of territoriality. We speculate that the formation of a conditioned place preference (CPP), an increased time allocation to the environment where a rewarding experience occurred, contributes to territoriality. Testosterone (T) plays an important role in modulating territorial behaviors and T pulses can induce a CPP. We confirmed previous findings in California mice (Peromyscus californicus) that T pulses can induce a CPP in singly-housed, but not group-housed males. Housing singly may be similar enough to dispersal in nature to initiate similar hormonal and neuroanatomical changes needed for the development of territoriality. We further revealed that T pulses interact with the single housing experience and appear to enhance the motivation to be aggressive towards a stimulus male. On a neural level, being singly housed upregulated levels of androgen receptors in the preoptic area, which positively correlated with the strength of the CPP. We speculate that this change in androgen sensitivity in the preoptic area is characteristic of males that have dispersed, making them more sensitive to T pulses. Also, single housing increased markers of synaptic plasticity in the nucleus accumbens, ventral and dorsal hippocampus, neural changes that may be associated with dispersal, reproduction and territory establishment. These behavioral and neural changes may reflect the life history transition from residing in the natal territory to dispersing and establishing a new territory.

An organism's ability to disperse influences many fundamental processes, from speciation and geographical range expansion to community assembly. However, the patterns and underlying drivers of variation in dispersal across species remain unclear, partly because standardised estimates of dispersal ability are rarely available. Here we present a global dataset of avian hand-wing index (HWI), an estimate of wing shape widely adopted as a proxy for dispersal ability in birds. We show that HWI is correlated with geography and ecology across 10,338 (>99%) species, increasing at higher latitudes and with migration, and decreasing with territoriality. After controlling for these effects, the strongest predictor of HWI is temperature variability (seasonality), with secondary effects of diet and habitat type. Finally, we also show that HWI is a strong predictor of geographical range size. Our analyses reveal a prominent latitudinal gradient in HWI shaped by a combination of environmental and behavioural factors, and also provide a global index of avian dispersal ability for use in community ecology, macroecology, and macroevolution.

Plasma testosterone can play an important role in promoting aggressive behaviors relating to territory defense in breeding male birds. Some birds defend territories also during the non-breeding phase, when testosterone circulates at basal levels. In such species, plasma levels of the pro-hormone dehydroepiandrosterone (DHEA) may support non-breeding territoriality by acting as a local substrate for sex steroids. To test this possible role of plasma DHEA, we examined the seasonal DHEA profile of male (and female) European nuthatches Sitta europaea: a male and female nuthatch pair will defend an all-purpose territory throughout the year. We hypothesized that plasma DHEA would be detectable in wintering nuthatches with a territory. However, only ca. half of the territorial wintering males (and females) displayed detectable DHEA levels, suggesting that plasma DHEA is not a major sex steroid precursor during non-breeding. Further, among hatching-year birds, plasma DHEA was significantly lower in territorial birds than in "floaters", i.e., subordinate birds without a territory. To experimentally examine the role of DHEA in non-breeding territoriality, we treated adult wintering males with DHEA and measured effects on aggressive responses to conspecific challenge. DHEA treatment elevated plasma levels of DHEA (and testosterone), but did not enhance territorial behaviors or their persistence. Taken together, our data suggest that DHEA (and, indeed, sex steroids per se) do not regulate non-breeding territoriality in the nuthatch. Given that territorial aggression in nuthatches is expressed year-round, a hormone for its activation may be redundant.

Year-around defense of extremely patchy habitat may require hormones that drive territorial behavior, but have no other costly physiological effects. The American pika (Ochotona princeps), but not the snowshoe hare (Lepus americanus) nor the eastern cottontail rabbit (Sylvilagus floridanus), exhibits this behavior. The former engages in contest competition against all individuals independent of sex to protect its territory in highly fragmented patches on mountain talus slopes; the latter in scramble competition in more continuous forests, grasslands, and shrublands. The hormonal basis for this difference in lagomorphs is unknown. Dehydroepiandrosterone is a prohormone produced by the zona reticularis of the adrenal cortex. It has no effect on aggressive behavior until converted in the brain to estrogen. We assessed levels (DHEA-S plus DHEA) in all species collected in the wild. In nonbreeding pikas, levels were 256 times higher than in hares and 22 times higher than in rabbits. Within species, females and males had similar levels. The proportion of the adrenal cortex devoted to the zona reticularis was significantly larger in pikas than in hares or rabbits. Our evidence is consistent with the hypothesis that dehydroepiandrosterone drives this individual-based, year-around territoriality in pikas. We propose a definitive experiment to determine this and recommend comparative studies in central Asia where there is high diversity of pika species whose behavior ranges from individual-based territoriality to colonial. Thus, we speculate that the wild American pika has the adrenal-brain nexus for all seasons and is an excellent model to understand how habitat drives the hormonal control of spacing behavior.

Female aggregation and male territoriality are considered to be hallmarks of polygynous mating systems. The development of genetic parentage assignment has called into question the accuracy of behavioral traits in predicting true mating systems. In this study we use 14 microsatellite markers to explore the mating system of one of the most behaviorally polygynous species, the California sea lion (Zalophus californianus). We sampled a total of 158 female-pup pairs and 99 territorial males across two breeding rookeries (San Jorge and Los Islotes) in the Gulf of California, Mexico. Fathers could be identified for 30% of pups sampled at San Jorge across three breeding seasons and 15% of sampled pups at Los Islotes across two breeding seasons. Analysis of paternal relatedness between the pups for which no fathers were identified (sampled over four breeding seasons at San Jorge and two at Los Islotes) revealed that few pups were likely to share a father. Thirty-one percent of the sampled males on San Jorge and 15% of the sampled males on Los Islotes were assigned at least one paternity. With one exception, no male was identified as the father of more than two pups. Furthermore, at Los Islotes rookery there were significantly fewer pups assigned paternity than expected given the pool of sampled males (p<0.0001). Overall, we found considerably lower variation in male reproductive success than expected in a species that exhibits behavior associated with strongly polygynous mating. Low variation in male reproductive success may result from heightened mobility among receptive females in the Gulf of California, which reduces the ability of males to monopolize groups of females. Our results raise important questions regarding the adaptive role of territoriality and the potential for alternative mating tactics in this species.

Many group-living species display strong sex biases in dispersal tendencies. However, gene flow mediated by apparently philopatric sex may still occur and potentially alters population structure. In our closest living evolutionary relatives, dispersal of adult males seems to be precluded by high levels of territoriality between males of different groups in chimpanzees, and has only been observed once in bonobos. Still, male-mediated gene flow might occur through rare events such as extra-group matings leading to extra-group paternity (EGP) and female secondary dispersal with offspring, but the extent of this gene flow has not yet been assessed.

Territoriality has been widely described across many animal taxa, where the acquisition and defence of a territory are critical for the fitness of an individual. Extensive evidence suggests that androgens are involved in the modulation of territorial behaviours in male vertebrates. Short-term increase of androgen following a territorial encounter appears to favour the outcome of a challenge. The "Challenge Hypothesis" proposed by Wingfield and colleagues outlines the existence of a positive feedback relationship between androgen and social challenges (e.g., territorial intrusions) in male vertebrates. Here we tested the challenge hypothesis in the highly territorial poison frog, Allobates femoralis, in its natural habitat by exposing males to simulated territorial intrusions in the form of acoustic playbacks. We quantified repeatedly androgen concentrations of individual males via a non-invasive water-borne sampling approach. Our results show that A. femoralis males exhibited a positive behavioural and androgenic response after being confronted to simulated territorial intrusions, providing support for the Challenge Hypothesis in a territorial frog.

Movements and habitat selection of predators shape ecological communities by determining the spatiotemporal distribution of predation risk. Although intraspecific interactions associated to territoriality and parental care are involved in predator habitat selection, few studies have addressed their effects simultaneously with those of prey and habitat distribution. Moreover, individuals require behavioural and temporal flexibility in their movement decisions to meet various motivations in a heterogeneous environment. To untangle the relative importance of ecological determinants of predator fine-scale habitat selection, we studied simultaneously several spatial, temporal, and behavioural predictors of habitat selection in territorial arctic foxes (Vulpes lagopus) living within a Greater snow goose (Anser caerulescens atlantica) colony during the reproductive season.