The social and defensive function of olfactory behaviors in a pair-living sexually monogamous primate.

Olfactory behaviors serve a wide variety of social functions in mammals. Odor may signal information about attributes of individuals important for mating and reproduction. Olfactory behaviors, such as scent-marking, may also function as part of home range or resource defense strategies. We assessed the potential social and home range defense functions of olfactory behavior in a pair-living and sexually monogamous primate, Azara's owl monkey (Aotus azarae), in the Argentinian Chaco. This is the most extensive investigation of owl monkey olfactory behaviors in the wild. Individuals regularly performed olfactory behaviors (group mean + SD = 1.3 + 0.5 per hour). The patterns were generally comparable to those observed in studies of captive owl monkeys, except that urine washing was the most common behavior in the wild, as opposed to scent-marking and genital inspections. Most olfactory behaviors were performed by adults, and there were striking sex differences in genital inspections: almost all consisted of an adult male inspecting the paired adult female. These findings suggest that olfactory behaviors play an important role in signaling and coordinating reproduction among owl monkeys, particularly during periods of female conception and pregnancy. Additionally, our research indicates that these behaviors may also serve as a defense strategy for maintaining the core area of their home ranges. This study offers the first assessment of the role of olfactory behaviors in reproductive contexts and home range defense in pair-living, monogamous platyrrhine primates.

Size matters: Natural experiments suggest the dear enemy effect is moderated by pack size in African wild dogs.

Remote monitoring of communal marking sites, or latrines, provides a unique opportunity to observe undisturbed scent marking behaviour of African wild dogs (Lycaon pictus). We used remote camera trap observations in a natural experiment to test behavioural scent mark responses to rivals (either familiar neighbours or unfamiliar strangers), to determine whether wild dogs exhibit the "dear enemy" or "nasty neighbour" response. Given that larger groups of wild dogs represent a greater threat to smaller groups, including for established residents, we predicted that the overarching categories "dear enemy" vs. "nasty neighbour" may be confounded by varying social statuses that exists between individual dyads interacting. Using the number of overmarks as a metric, results revealed an interaction between sender and receiver group size irrespective of familiarity consistent with this prediction: in general, individuals from large resident packs overmarked large groups more than they overmarked smaller groups, whereas individuals from smaller packs avoided overmarking larger groups, possibly to avoid detection. Monitoring a natural system highlights variables such as pack size that may be either overlooked or controlled during scent presentation experiments, influencing our ability to gain insights into the factors determining territorial responses to rivals.

Mesocarnivores vary in their spatiotemporal avoidance strategies at communications hubs of an apex carnivore.

Mesocarnivores face interspecific competition and risk intraguild predation when sharing resources with apex carnivores. Within a landscape, carnivores across trophic levels may use the same communication hubs, which provide a mix of risks (injury/death) and rewards (gaining information) for subordinate species. We predicted that mesocarnivores would employ different strategies to avoid apex carnivores at shared communication hubs, depending on their trophic position. To test our prediction, we examined how different subordinate carnivore species in the Santa Cruz Mountains of California, USA, manage spatial overlap with pumas (Puma concolor), both at communication hubs and across a landscape-level camera trap array. We estimated species-specific occurrence, visitation rates, temporal overlap, and Avoidance-Attraction Ratios from camera traps and tested for differences between the two types of sites. We found that mesocarnivores generally avoided pumas at communication hubs, and this became more pronounced when pumas scent-marked during their most recent visit. Coyotes (Canis latrans), the pumas' closest subordinate competitor in our system, exhibited the strongest avoidance at communication hubs. Gray foxes (Urocyon cinereoargenteus) avoided pumas the least, which may suggest possible benefits from pumas suppressing coyotes. Overall, mesocarnivores exhibited various spatiotemporal avoidance strategies at communication hubs rather than outright avoidance, likely because they benefit from information gained while 'eavesdropping' on puma activity. Variability in avoidance strategies may be due to differential predation risks, as apex carnivores often interact more aggressively with their closest competitors. Combined, our results show how apex carnivores trigger complex species interactions across the entire carnivore guild and how trophic position determines behavioral responses and subsequent space use of subordinate mesocarnivores across the landscape.

Male dingo urinary scents code for age class and wild dingoes respond to this information.

Chemical information in canid urine has been implicated in territoriality and influences the spacing of individuals. We identified the key volatile organic compound (VOC) components in dingo (Canis lupus dingo) urine and investigated the potential role of scents in territorial spacing. VOC analysis, using headspace gas chromatography-mass spectrometry (GC-MS), demonstrated that the information in fresh urine from adult male dingoes was sufficient to allow statistical classification into age categories. Discriminant function analyses demonstrated that the relative amounts or combinations of key VOCs from pre-prime (3-4 years), prime (5-9 years), and post-prime (≥10 years) males varied between these age categories, and that scents exposed to the environment for 4 (but not 33) days could still be classified to age categories. Further, a field experiment showed that dingoes spent less time in the vicinity of prime male dingo scents than other scents. Collectively, these results indicate that age-related scent differences may be discriminable by dingoes. Previous authors have suggested the potential to use scent as a management tool for wild canids by creating an artificial territorial boundary/barrier. Our results suggest that identifying the specific signals in prime-age male scents could facilitate the development of scent-based tools for non-lethal management.

Effects of intrauterine position during gestation on specific endocrine and behavioral parameters that impact reproduction in domestic rabbits.

Prior studies from others, performed in a different breed, reported that doe rabbits developing between two male siblings (2 M) during gestation display characteristics indicative of masculinization: larger anogenital distance (AGD), larger submandibular glands, and higher chinning frequency than females with zero (0 M) or one (1 M) contiguous brothers. Similar effects are provoked by injecting androgens to the pregnant doe suggesting that prenatal androgen exposure masculinizes female embryos. To further understand the scope of such masculinization we compared 0 M, 1 M, and 2 M females regarding behavioral, neuroendocrine, and somatic parameters, related or not to reproduction. IUP did not impact: body weight, sexual receptivity, mating-induced LH secretion, maternal nest-building, litter size, or milk output. At puberty: a) chinning frequency was: 0 M and males>1 M and 2 M; b) ambulation in open field was lowest in 1 M females and males. IUP effects on AGD were significant only on postnatal day 1: 0 M, 1 M, and males>2 M, in contrast to earlier study. Willingness to nurse at delivery was less frequent in 2 M than in 1 M and 0 M does and correlated with nursing occurrence across lactation. Does that did not nurse at parturition delivered fewer kits/min than those that nursed then, regardless of IUP. The duration of nursing bouts across lactation was significantly longer in the1 M and 2 M does that showed this behavior on postpartum days 1-20. Our findings indicate that IUP is associated with alterations in specific aspects of postpartum maternal behavior.

Latrine site selection by African clawless otters, Aonyx capensis, and their behavior during latrine visitations.

Latrine sites are used as areas for the deposition of scent-containing excretions and play important roles in intraspecific olfactory communication, territoriality, sexual attraction, and defense behaviors of many mammals. African clawless otters (Aonyx capensis) likely use latrine sites as primary areas for scent marking and scent communication but no studies to date have investigated their potential role or site selection. We assessed latrine site selection at 2 spatial scales (micro- and macroscale) and recorded behaviors via camera trap recordings. Thirty-eight latrine sites were identified and assessed at 2 locations in Mtunzini on the north coast of KwaZulu-Natal, South Africa (uMlalazi Nature Reserve and Zini Fish Farm) during the months of August to November 2021. Latrine sites were identified through several intensive surveys, while we characterized nonselected sites through a systematic sampling approach. Latrine and control sites were inventoried along a 52-m buffer around all water bodies in both study areas. At each site we measured a series of potential environmental predictors, including horizontal and vertical vegetation cover, surface slope, and averaged wind speeds for days classified as relatively wind-still and relatively windy. To assess the relative role of various environmental predictors, we used a binomial generalized linear model resource selection function to model both spatial scales of latrine site selection. The majority of latrine sites were located at the ecotone between 2 vegetation units or between a vegetation unit and a water source. At a macroscale, latrine sites were associated with areas containing little vegetative substrate cover and minimal canopy cover. The top-ranked models at the microscale also indicated that latrine sites were characterized as occurring in open areas with less canopy and horizontal cover and on flatter areas that are relatively protected against wind. The most common behaviors recorded at 3 latrine sites were the "jiggle dance" (42%) and sniffing (29%). We hypothesize that otters evaluate numerous environmental parameters to enhance the functionality of latrine sites. For example, sites with little vegetative cover may increase the conspicuousness of latrines to conspecifics, while areas exposed to less wind likely aid in the retention of scent. Ongoing research is characterizing the behaviors of otters around latrines and chemical signatures of latrine sites in an effort to facilitate interpretation of their social function to African clawless otters.

Signal detection theory applied to giant pandas: Do pandas go out of their way to make sure their scent marks are found?

Inter-animal communication allows signals released by an animal to be perceived by others. Scent-marking is the primary mode of such communication in giant pandas (Ailuropoda melanoleuca). Signal detection theory propounds that animals choose the substrate and location of their scent marks so that the signals released are transmitted more widely and last longer. We believe that pandas trade-off scent-marking because they are an energetically marginal species and it is costly to generate and mark chemical signals. Existing studies only indicate where pandas mark more frequently, but their selection preferences remain unknown. This study investigates whether the marking behavior of pandas is consistent with signal detection theory. Feces count, reflecting habitat use intensity, was combined with mark count to determine the selection preference for marking. The results showed that pandas preferred to mark ridges with animal trails and that most marked tree species were locally dominant. In addition, marked plots and species were selected for lower energy consumption and a higher chance of being detected. Over 90% of the marks used were the longest-surviving anogenital gland secretion marks, and over 80% of the marks were oriented toward animal trails. Our research demonstrates that pandas go out of their way to make sure their marks are found. This study not only sheds light on the mechanisms of scent-marking by pandas but also guides us toward more precise conservation of the panda habitat.

Scent mark signal investment predicts fight dynamics in house mice.

Signals mediate competitive interactions by allowing rival assessment, yet are often energetically expensive to produce. One of the key mechanisms maintaining signal reliability is social costs. While the social costs of over-signalling are well known, the social costs of under-signalling are underexplored, particularly for dynamic signals. In this study, we investigate a dynamic and olfactory-mediated signalling system that is ubiquitous among mammals: scent marking. Male house mice territorially scent mark their environment with metabolically costly urine marks. Competitive male mice are thought to deposit abundant scent marks in the environment. However, we recently identified a cohort of low-marking males that win fights. We hypothesized that there may be social costs imposed on individuals who under-invest in signalling. Here we find that scent mark investment predicts fight dynamics. Winning males that produce fewer scent marks prior to a fight engage in more intense fights that take longer to resolve. This effect appears to be driven by an unwillingness among losers to acquiesce to weakly signalling winners. We, therefore, find evidence for rival assessment of scent marks as well as social costs to under-signalling. This supports existing hypotheses for the importance of social punishment in maintaining optimal signalling equilibria. Our results further highlight the possibility of diverse signalling strategies in house mice.

Reading the signs: Camera-trapping provides new insights on scent marking in the large-antlered muntjac (Muntiacus vuquangensis).

We present evidence of scent marking in the large-antlered muntjac (Muntiacus vuquangensis). Given the importance of scent marking in individual recognition among ungulates, this behavior may serve to communicate the fitness cost of antagonistic interactions among rival males and could serve as a mechanism for mate assessment among females.

Faded neural projection from the posterior bed nucleus of the stria terminalis to the lateral habenula contributes to social signaling deficit in male BTBR mice as a mouse model of autism.

BTBR T+ Itpr3tf/J (BTBR) mice display several behavioral characteristics, including social deficits resembling the core symptoms of human autism. Atypical social behaviors include sequential processes of assembled cognitive-behavior components, such as recognition, investigatory assessment, and signaling response. This study aimed to elucidate the neural circuits responsible for the regulation of the social signaling response, as shown by scent marking behavior in male mice. We first assessed the recognition and investigatory patterns of male BTBR mice compared to those of C57BL/6 J (B6) mice. Next, we examined their scent-marking behavior as innate social signaling responses adjusted to a confronted feature of social stimuli and situations, along with the expression of c-Fos as a marker of neuronal activity in selected brain areas involved in the regulation of social behavior. The function of the targeted brain area was confirmed by chemogenetic manipulation. We also examined the social peptides, oxytocin and vasopressin neurons of the major brain regions that are associated with the regulation of social behavior. Our data indicate that male BTBR mice are less responsive to the presentation of social stimuli and the expression of social signaling responses, which is paralleled by blunted c-Fos responsivity and vasopressin neurons morphological changes in selected brain areas, including the posterior bed nucleus of the stria terminalis (pBnST) and lateral habenula (LHb) in BTBR mice. Further investigation of LHb function revealed that chemogenetic inhibition and activation of LHb activity can induce a change in scent marking responses in both B6 and BTBR mice. Our elucidation of the downstream LHb circuits controlling scent marking behavior indicates intact function in BTBR mice. The altered morphological characteristics of oxytocin neurons in the paraventricular nucleus of the hypothalamus and vasopressin-positive neurons and axonal projections in the pBnST and LHb appear to underlie the dysfunction of scent marking responses in BTBR mice. (300/300 words).

Effects of a GnRH agonist on sex behavior in females of the southern giant pouched rat.

In general, males should be particularly attentive to cues of sexual availability and females should advertise accordingly. Vaginal patency (i.e., the openness of the vagina) is a reliable indicator of sexual maturity; if the vagina is closed, the female is unable to copulate. The southern giant pouched rat (Cricetomys ansorgei) is unusual because females can have fully fused vaginal openings (i.e., vaginal nonpatency) despite being considered 'adults' by other metrics. Moreover, some females reversibly close their vaginal openings. Thus, vaginal patency in the pouched rat is a 'flexible' reproductive state. We subcutaneously implanted a long-acting GnRH agonist (deslorelin), which over time inhibits sex steroid secretion, to better understand the endocrinology and social behavior relating to vaginal patency in this species. We hypothesized that altering GnRH would impact both patency and behavior through its effects on circulating levels of estradiol. Six months of deslorelin treatment did not alter vaginal patency. Behaviorally, deslorelin-treated females spent less time interacting with, and were more aggressive towards males (compared to controls). Notably, deslorelin did not alter female scent marking. We conclude that behavioral receptivity, but not vaginal patency, is impacted by GnRH hormonal cascades in the pouched rat.

Is scat marking a reliable tool for otter census and surveys at the landscape scale?

Biological significance of scat marking by otters has been a controversial subject among scientists. Using multiyear (2014-2017) data of otter spraint counts in South Korea, this study aimed to test whether the observed pattern of spraint presence/absence is driven by detection error and if/how scat counts can be a proxy for otter abundance at the landscape scale. To test the first hypothesis, spraint presence/absence was analyzed through occupancy models, which relied on environmental variables related to otter detectability and presence. Spraint count models were used to test the second hypothesis against resource-related covariates in combination with landscape, anthropogenic, and climate variables through machine learning algorithms (MLAs). The detection probability has specifically decreased in areas characterized by high rainfall and human population densities, whereas the probability has increased near food-rich sites, characterized by high marking frequencies. The temporal trends of spraint count predictions were in line with changes in the diversity of fish communities in 2014-2017 instead of fish biomass, suggesting that the availability of feeding resources is higher where fish communities are more diverse. Because diverse fish communities can attract otters, fish diversity conservation is critical for preserving this mammal's populations. This fine scale four-year monitoring has contributed to the disentanglement of the role of spraint presence/absence and spraint counts in detectability and population trends. This will assist in identifying key resource areas and planning strategies to promote otter conservation and dispersal dynamics.

Do male meadow voles (Microtus pennsylvanicus) always prefer and signal to 'popular' females? The role of social information context.

For males, the behaviour of rival conspecifics can provide vital information about finding reproductively available females, and which have high-quality phenotypes. While valuable, this information represents sperm competition risk and even the potential for rejection by females. Thus, males should show flexibility in signalling behaviours towards females based on social information acquired with emphasis on the context of that information. We examined male meadow vole scent-marking behaviours and preferences for females based on previously obtained social information across different contexts. Social information context resulted in increased scent marking depending on the social odour pairing. Males scent marked more near a female whose odour had associated with three older rivals than one younger rival. But males marked equally toward a female whose odour had associated with three younger rivals and a female whose odour had associated with one older rival. This demonstrates that social odours are not of all equal value and that males can distinguish differences. Males then use these differences to tailor their signalling towards potentially high-quality phenotype females thereby maximizing their reproductive opportunities.

Linking the Morphology of Sternal Glands to Rubbing Behavior by Vespa soror (Hymenoptera: Vespidae) Workers During Recruitment for Group Predation.

The activities of social insect colonies are supported by exocrine glands and the tremendous functional diversity of the compounds that they secrete. Many social wasps in the subfamilies Vespinae and Polistinae have two sternal glands-the van der Vecht and Richards' glands-that vary in their features and function across the species in which they are found. Field observations suggest that giant hornets use secretions from the van der Vecht gland to chemically mark targeted nests when workers initiate group attacks on social insect prey. However, descriptions of giant hornets' sternal glands and details about their recruitment behavior are lacking. We describe the morphology of the sternal glands of the giant hornet Vespa soror du Buysson and consider their potential to contribute to a marking pheromone. We also assess the gastral rubbing behavior of workers as they attacked Apis cerana F. (Hymenoptera: Apidae) colonies. V. soror workers have well-developed van der Vecht and Richards' glands on their terminal gastral sternites, with morphologies that robustly support the synthesis, storage, and dissemination of their secretory products. Observations confirm that the van der Vecht gland is exposed during gastral rubbing, but that the Richards' gland and glands associated with the sting apparatus may also contribute to a marking pheromone. Workers briefly but repeatedly rubbed their gasters around hive entrances and on overhead vegetation. Colonies were heavily marked over consecutive attacks. Our findings provide insight into the use of exocrine secretions by giant hornets as they recruit nestmates to prey colonies for group attacks.

Exocrine scent marking: Coordinative role of arginine vasopressin in the systemic regulation of social signaling behaviors.

Arginine vasopressin (AVP) is a neurohypophysial hormone that coordinatively regulates central socio-emotional behavior and peripheral control of antidiuretic fluid homeostasis. Most mammals, including rodents, utilize exocrine or urine-contained scent marking as a social signaling tool that facilitates social adaptation. The exocrine scent marking behavior is postulated to fine-tune sensory and cognitive abilities to recognize key social features via exocrine/urinary olfactory cues and subsequently control exocrine deposition or urinary marking through the mediation of osmotic fluid balance. AVP is implicated as a major player in controlling both recognition and signaling responses. This review provides constructive hypotheses on the coordinative processes of the AVP neurohypophysial circuits in the systemic regulations of fluid control and social-communicative behavior, via the expression of exocrine scent marking, and further emphasizes a potential role of AVP in a common mechanism underlying social communication in rodents.

Contrasting central and systemic effects of arginine-vasopressin on urinary marking behavior as a social signal in male mice.

Arginine-vasopressin (AVP) is a neurohypophyseal peptide that plays a critical role in the regulation of social behavior in mammals. Neuronal AVP regulates male-specific social signaling processes, such as exocrine urinary scent deposition and marking behavior in mice. In the periphery, AVP is transported to the portal bloodstream and acts as an antidiuretic hormone. These AVP dynamics imply that the central role of AVP in the stimulation of urinary marking is dissociated with the peripheral role of AVP in the retention of osmotic conditions. Using male BALB/c mice as subjects, peripheral injection of AVP decreased urinary marking and urination. In contrast, a central infusion of AVP facilitated urinary marking with no effect on urination, while an antagonist of the AVP 1a receptor inhibited marking. Centrally AVP-injected mice also exhibited typical behaviors, such as hiccough/sneeze-like reactions and flash scratching, particularly when confronted with a stimulus mouse through a wire mesh screen. Significant expression of these typical reactions in these mice resulted in the disruption of marking deposition. Further analysis of AVP synthesis illustrated that AVP levels increased in the midbrain but not in the circulation immediately after the test, particularly when confronted with a stimulus mouse. The central AVP regulates urinary marking and other typical behaviors in a dose- and situation-dependent manner. The sequential process implies that centrally synthesized AVP may be secreted into the circulation following immediate neuronal processes, and then peripheral AVP acts as an antidiuretic hormone on urinary marking behavior.

Effect of proximity to females on integument damage caused by self-enurination in male goats.

In male goats, self-enurination (SE) is the downward turning of the head and shoulders while urinating onto the face and front legs. Although it provides important chemical cues to females, other males, and even self, it is a costly behavior that can create a range of problems including erythema, irritation, hair loss, and compromised skin. It was hypothesized that the extent of integument damage from SE on bucks' faces and front legs would be increased by housing bucks near females. Four bucks were housed with fence-line contact to females ("Near" bucks), and four bucks were housed without fence-line contact to females ("Far" bucks). Each buck was photographed every other week over an 18-wk period during the breeding season. During each imaging session, seven different photographic views were captured, and burn areas were quantified for each buck. Overall, more Near bucks had urine burn than Far bucks, and Near bucks developed urine burn earlier in the breeding season than did Far bucks. Housing bucks close to females increases the extent of integument damage from SE. These findings may help goat breeders develop management practices to improve animal well-being by minimizing urine burn injury to bucks.

Sexual Dimorphism in Crowned Lemur Scent-Marking.

Primates are traditionally considered to have a poor sense of smell. However, olfaction is important for non-human primates as demonstrated by conspicuous scent-marking behaviours in lemurs. We studied two pairs (n = 4) of crowned lemurs (Eulemur coronatus) housed at Colchester and Twycross zoos (UK) by combining behavioural observations and chemical analyses of scent-marks and glandular swabs. We recorded observations of olfactory behaviours for 201 h using instantaneous scan sampling. We investigated the volatile compounds of ano-genital odour secretions (n = 16) using solid-phase microextraction and gas chromatography-mass spectrometry. Males scent-marked most frequently, displaying ano-genital marking for allomarking, head marking for countermarking and wrist marking in specific areas of the enclosure. Females displayed ano-genital marking, predominantly on feeding devices. We detected a total of 38 volatile components in all male ano-genital scent-marks and 26 in all female samples of ano-genital odour secretions, including a series of esters, aldehydes, ketones, alcohols, terpenes, volatile fatty acids and hydrocarbons that have been identified in odour profiles of other primates. In conclusion, we found sexual dimorphism in crowned lemur scent-marking. Male head and wrist marking behaviours might play defensive territorial functions, while ano-genital marking would be related to socio-sexual communication as chemical mate-guarding. Female ano-genital marking might be involved in resource defense.

On the trail of primate scent signals: A field analysis of callitrichid scent-gland secretions by portable gas chromatography-mass spectrometry.

Chemosignals are mediators of social interactions in mammals, providing con- and hetero-specifics with information on fixed (e.g., species, sex, group, and individual identity) and variable (e.g., social, reproductive, and health status) features of the signaler. Yet, methodological difficulties of recording and quantifying odor signals, especially in field conditions, have hampered studies of natural systems. We present the first use of the Torion® portable gas chromatography-mass spectrometry (GC-MS) instrument for in situ chemical analysis of primate scents. We collected and analyzed swab samples from the scent glands and skin from 13 groups (57 individuals) of two sympatric species of wild emperor tamarins, Saguinus imperator, and Weddell's saddleback tamarins, Leontocebus weddelli (Callitrichidae). In total, 11 compounds of interest (i.e., probably derived from the animals) could be detected in the samples, with 31 of 215 samples containing at least one compound of interest. The composition of these 31 samples varied systematically with species, group, sex, and breeding status. Moreover, we tentatively identified seven of the compounds of interest as methyl hexanoate, benzaldehyde, ethyl hexanoate, acetophenone, a branched C15 alkane, 4-methoxybenzaldehyde, and hexadecan-1-ol. As the field of primate semiochemistry continues to grow, we believe that portable GC-MS instruments have the potential to help make progress in the study of primate chemosignaling in field conditions, despite limitations that we encountered. We further provide recommendations for future use of the Torion® portable GC-MS for in situ analyses.

Brown bear communication hubs: patterns and correlates of tree rubbing and pedal marking at a long-term marking site.

Chemical communication is important for many species of mammals. Male brown bears, Ursus arctos, mark trees with a secretion from glands located on their back. The recent discovery of pedal glands and pedal-marking at a site used for tree-rubbing led us to hypothesize that both types of marking form part of a more complex communication system. We describe the patterns of chemical communication used by different age and sex classes, including differences in the roles of these classes as information providers or receivers over four years at a long-term marking site. Using video recordings from a camera trap, we registered a total of 285 bear-visits and 419 behavioral events associated with chemical communication. Bears visited the site more frequently during the mating season, during which communication behaviors were more frequent. A typical visit by male bears consisted of sniffing the depressions where animals pedal mark, performing pedal-marking, sniffing the tree, and, finally, rubbing against the trunk of the tree. Adult males performed most pedal- and tree-marking (95% and 66% of the cases, respectively). Males pedal-marked and tree-rubbed in 81% and 48% of their visits and sniffed the pedal marks and the tree in 23% and 59% of visits, respectively. Adult females never pedal marked, and juveniles did so at very low frequencies. Females rubbed against the tree in just 9% of their visits; they sniffed the tree and the pedal marks in 51% and 21% of their visits, respectively. All sex and age classes performed pedal- and tree-sniffing. There were significant associations between behaviors indicating that different behaviors tended to occur during the same visit and were more likely if another individual had recently visited. These associations leading to repeated marking of the site can promote the establishment of long-term marking sites. Marking sites defined by trees and the trails leading to them seem to act as communication hubs that brown bears use to share and obtain important information at population level.