Age-sex Classes Research Articles

Camera Traps Reveal Variation in Diel Activity Patterns among Different Sex-Age Classes in Brown Bears in Hokkaido

Camera Traps Reveal Variation in Diel Activity Patterns among Different Sex-Age Classes in Brown Bears in Hokkaido

Protection service of a leading silverback male from external threats in wild western gorillas.

Primate males normally protect reproductive females, genetic offspring, and other relatives from external threats. Nevertheless, male protection of group members other than the above individuals is widely reported. Here, we show qualitative data on a silverback's charging behaviors toward human observers (predator surrogates) to protect group members having various age-sex and kinship traits in a group of wild western gorillas containing one reproductive male. We observed 106 and 33 charging behaviors by the leading silverback in two separate study periods. Two natal infants were often involved in his protective charging. Further, the silverback provided protection services to reproductive females. Surprisingly, immigrant individuals (i.e., unrelated to the silverback), including a wide range of age-sex classes, were also protected multiple times. His protection services for natal infants and adult females can be interpreted as a form of parenting effort and mating effort, respectively. Further, those for some immigrant immatures accompanied by their mothers can be considered part of mating effort, advertising his quality as a mate to the mothers. Finally, his charging behaviors to protect immigrant young males, who could be reproductive threats to him, may be due to group augmentation benefits. That is, the recruitment of additional males in exchange for protection services would improve the ability of group defense. Protection services of the leading silverback in the one-male group of western gorillas, in which members of various age-sex classes and kinship traits coexist, could be interpreted by some existing functional explanations.

Enriched social environment affects the utterance of acoustic signals of captive François' langurs

Vocal communication serves as an important mode of interaction among animals. The quantitative analysis of acoustic signals holds scientific value in highlighting the importance of understanding the social-ecological contexts and their effects on acoustic communication. The results can provide a direct scientific guidance for enhancing the welfare and management of captive animals. From January to February 2022 and from December 2022 to February 2023, we conducted a study involving 30 captive François' langurs (Trachypithecus francoisi) housed in nine enclosures. We collected acoustic signals along with their associated social-ecological contexts and recorded the social composition and number of François' langurs in each enclosure. We in total identified 23 call types based on human ear auditory, spectrogram visual inspection, and quantitative analysis of 37 acoustic parameters. Our findings revealed that adult males exhibited nine call types (five adult male-specific types), adult females 11 call types (seven adult female-specific types), juveniles eight call types (two juvenile-specific types), and infants five call types (two infant-specific types). The asymmetry in call types indicated distinct social roles among different sex-age classes of François' langurs. We found that a complex social composition or larger group size resulted in a more diverse range of call types and increased utterance frequency among François' langurs. The increased utterance frequency was linked to calls used in neutral (contact), affiliation, mating, and vigilance contexts, indicating that a diverse social setting encourages social interaction and boosts the occurrence of natural behaviors, and may also result in increased tension and anxiety. Implementing a one-male multi-female model with ample breeding space for captive François' langurs was identified as beneficial for enhancing vocal communication and improving animal welfare. This study established the first vocal repertoire of François' langurs, offering a novel scientific approach to evaluating the welfare of this endangered species through vocal behaviors.

Urbanization does not increase "object curiosity" in vervet monkeys, but semi-urban individuals selectively explore food-related anthropogenic items.

Urban environments expose animals to abundant anthropogenic materials and foods that facilitate foraging innovations in species with opportunistic diets and high behavioral flexibility. Neophilia and exploration tendency are believed to be important behavioral traits for animals thriving in urban environments. Vervet monkeys (Chlorocebus pygerythrus) are one of few primate species that have successfully adapted to urban environments, thus making them an ideal species to study these traits. Using a within-species cross-habitat approach, we compared neophilia and exploration of novel objects (jointly referred to as "object curiosity") between semi-urban, wild, and captive monkeys to shed light on the cognitive traits facilitating urban living. To measure "object curiosity," we exposed monkeys to various types of novel stimuli and compared their approaches and explorative behavior. Our results revealed differences in the number of approaches and explorative behavior toward novel stimuli between the habitat types considered. Captive vervet monkeys were significantly more explorative than both semi- urban and wild troops, suggesting that positive experiences with humans and lack of predation, rather than exposure to human materials per se, influence object curiosity. Across habitats, juvenile males were the most explorative age-sex class. This is likely due to males being the dispersing sex and juveniles being more motivated to learn about their environment. Additionally, we found that items potentially associated with human food, elicited stronger explorative responses in semi-urban monkeys than non-food related objects, suggesting that their motivation to explore might be driven by "anthrophilia", that is, their experience of rewarding foraging on similar anthropogenic food sources. We conclude that varying levels of exposure to humans, predation and pre-exposure to human food packaging explain variation in "object curiosity" in our sample of vervet monkeys.

Ecology of fear alters behavior of grizzly bears exposed to bear-viewing ecotourism.

Humans are perceived as predators by many species and may generate landscapes of fear, influencing spatiotemporal activity of wildlife. Additionally, wildlife might seek out human activity when faced with predation risks (human shield hypothesis). We used the anthropause, a decrease in human activity resulting from the COVID-19 pandemic, to test ecology of fear and human shield hypotheses and quantify the effects of bear-viewing ecotourism on grizzly bear (Ursus arctos) activity. We deployed camera traps in the Khutze watershed in Kitasoo Xai'xais Territory in the absence of humans in 2020 and with experimental treatments of variable human activity when ecotourism resumed in 2021. Daily bear detection rates decreased with more people present and increased with days since people were present. Human activity was also associated with more bear detections at forested sheltered sites and less at exposed sites, likely due to the influence of habitat on bear perception of safety. The number of people negatively influenced adult male detection rates, but we found no influence on female with young detections, providing no evidence that females responded behaviorally to a human shield effect from reduced male activity. We also observed apparent trade-offs of risk avoidance and foraging. When salmon levels were moderate to high, detected bears were more likely to be females with young than adult males on days with more people present. Should managers want to minimize human impacts on bear activity and maintain baseline age-sex class composition at ecotourism sites, multiday closures and daily occupancy limits may be effective. More broadly, this work revealed that antipredator responses can vary with intensity of risk cues, habitat structure, and forage trade-offs and manifest as altered age-sex class composition of individuals using human-influenced areas, highlighting that wildlife avoid people across multiple spatiotemporal scales.

Call combination in African forest elephants Loxodonta cyclotis.

Syntax, the combination of meaning-devoid phonemes into meaningful words, which in turn are combined in structurally and semantically complex sentences, is fundamental to the unlimited expressiveness of human languages. Studying the functions of call combinations in non-human species provides insights into the evolution of such syntactic capabilities. Here, we investigated the combination of high amplitude broadband calls with low frequency rumble vocalizations in a highly social species, the African forest elephant Loxodonta cyclotis. Rumbles play an integral role in coordinating social interactions by transmitting socially relevant information, including individual identity. By contrast, broadband calls, such as roars, are thought to function as signals of distress and urgency as they are typically produced in situations of high emotional intensity. Functional changes associated with the combination of these calls remain little understood. We found that call combinations were produced by all age-sex classes but were most prevalent in immature individuals. We found that rumbles used singularly occurred in all five investigated social contexts, whereas single broadband calls were restricted to two resource-related contexts. Call combinations also occurred in all five contexts, suggesting an increase in the functional use of broadband calls when combined with rumbles, analogous to the generativity brought about through syntax in human speech. Moreover, combining calls appeared to lead to functional shifts towards high-stake contexts. Call combinations were more likely in competition contexts compared to single rumbles, and more likely in separation contexts compared to single broadband calls. We suggest that call combination in forest elephants may aide to reduce message ambiguity in high-stake situation by simultaneously communicating distress and individual identity, which may be critical to secure access to resources, reduce the risk of injury and to reunite with or recruit the support of the family group.

Satellite tracking of American Woodcock reveals a gradient of migration strategies

Abstract Diversity in behavior is important for migratory birds in adapting to dynamic environmental and habitat conditions and responding to global change. Migratory behavior can be described by a variety of factors that comprise migration strategies. We characterized variation in migration strategies in American Woodcock (Scolopax minor), a migratory gamebird experiencing long-term population decline, using GPS data from ~300 individuals tracked throughout eastern North America. We classified woodcock migratory movements using a step-length threshold, and calculated characteristics of migration related to distance, path, and stopping events. We then used principal components analysis (PCA) to ordinate variation in migration characteristics along axes that explained different fundamental aspects of migration, and tested effects of body condition, age-sex class, and starting and ending location on PCA results. The PCA did not show evidence for clustering, suggesting a lack of discrete strategies among groups of individuals; rather, woodcock migration strategies existed along continuous gradients driven most heavily by metrics associated with migration distance and duration, departure timing, and stopping behavior. Body condition did not explain variation in migration strategy during the fall or spring, but during spring adult males and young females differed in some characteristics related to migration distance and duration. Starting and ending latitude and longitude, particularly the northernmost point of migration, explained up to 61% of the variation in any one axis of migration strategy. Our results reveal gradients in migration behavior of woodcock, and this variability should increase the resilience of woodcock to future anthropogenic landscape and climate change.

Spring haul-out behavior of seals in the Bering and Chukchi Seas: implications for abundance estimation.

Ice-associated seals rely on sea ice for a variety of activities, including pupping, breeding, molting, and resting. In the Arctic, many of these activities occur in spring (April through June) as sea ice begins to melt and retreat northward. Rapid acceleration of climate change in Arctic ecosystems is therefore of concern as the quantity and quality of suitable habitat is forecast to decrease. Robust estimates of seal population abundance are needed to properly monitor the impacts of these changes over time. Aerial surveys of seals on ice are an efficient method for counting seals but must be paired with estimates of the proportion of seals out of the water to derive population abundance. In this paper, we use hourly percent-dry data from satellite-linked bio-loggers deployed between 2005 and 2021 to quantify the proportion of seals hauled out on ice. This information is needed to accurately estimate abundance from aerial survey counts of ice-associated seals (i.e., to correct for the proportion of animals that are in the water, and so are not counted, while surveys are conducted). In addition to providing essential data for survey 'availability' calculations, our analysis also provides insights into the seasonal timing and environmental factors affecting haul-out behavior by ice-associated seals. We specifically focused on bearded (Erignathus barbatus), ribbon (Histriophoca fasciata), and spotted seals (Phoca largha) in the Bering and Chukchi seas. Because ringed seals (Phoca (pusa) hispida) can be out of the water but hidden from view in snow lairs analysis of their 'availability' to surveys requires special consideration; therefore, they were not included in this analysis. Using generalized linear mixed pseudo-models to properly account for temporal autocorrelation, we fit models with covariates of interest (e.g., day-of-year, solar hour, age and sex class, wind speed, barometric pressure, temperature, precipitation) to examine their ability to explain variation in haul-out probability. We found evidence for strong diel and within-season patterns in haul-out behavior, as well as strong weather effects (particularly wind and temperature). In general, seals were more likely to haul out on ice in the middle of the day and when wind speed was low and temperatures were higher. Haul-out probability increased through March and April, peaking in May and early June before declining again. The timing and frequency of haul-out events also varied based on species and age-sex class. For ribbon and spotted seals, models with year effects were highly supported, indicating that the timing and magnitude of haul-out behavior varied among years. However, we did not find broad evidence that haul-out timing was linked to annual sea-ice extent. Our analysis emphasizes the importance of accounting for seasonal and temporal variation in haul-out behavior, as well as associated environmental covariates, when interpreting the number of seals counted in aerial surveys.

Activity budget of a group of black-horned capuchin monkeys (Sapajus nigritus) in an urban environment

Urban growth and human impacts on the environment have forced animals to adjust to habitat fragmentation and reduced home ranges. Capuchin monkeys are known for their great social and behavioral flexibility, occupying even highly urbanized environments in a way that the time budget of this primate in synanthropic situation may be affected by the area they inhabit. This study aims to analyze the activity budget of a group of Sapajus nigritus living in an anthropized area, 1) comparing the behavioral frequencies in urbanized areas and forest fragments; 2) comparing behavioral frequencies in different sex-age classes. During the study, the number of individuals ranged from 35 to 40 individuals identified based on sex-age classes. Behavioral data were collected using the instantaneous scan sampling method, for two minutes with eight-minute intervals. We obtained 319 scans over 28 days, distributed between November 2021 and June 2022, with eight hours per day. We compared the behaviors different areas and between sex-age classes using the Kruskal-Wallis’s test. Overall, the group performed a higher frequency of traveling (21.22%), followed by foraging (18.07%), feeding (16.57%) and vigilance (15.61%). The frequency of behaviors varied between areas, with vigilance, social, resting, interaction with humans and self-activity more frequent in urbanized areas compared to forest fragments. We also found variation between the sex-age classes, primarily with juveniles foraging more and adults performing more vigilance. The differences in the behaviors performed by the group express the behavioral flexibility of S. nigritus, adapting its activity pattern according to the area occupied.

Attributing Ethiopian animal health losses to high-level causes using expert elicitation

The Global Burden of Animal Diseases programme is currently working to estimate the burden of animal health loss in Ethiopia. As part of this work, structured expert elicitation has been trialled to attribute the proportion of animal health losses due to three independent and exhaustive high-level causes (infectious, non-infectious, and external). Separate in-person workshops were conducted with eight cattle, nine small ruminant, and eight chicken experts. Following the Investigate-Discuss-Estimate-Aggregate protocol for structured expert elicitation, estimates were obtained for the proportion of animal health loss due to high-level causes in different combinations of health loss, species, age-sex class, and production system. Three-point questions were used to inform beta-pert distributions and capture uncertainty in estimates. Individual expert estimates were aggregated by quantile mean to produce average distributions. Random samples from these average distributions estimated that infectious causes inflict the highest proportion of health loss in Ethiopia, with at least 40 % of health losses estimated to be due to infectious causes in all categories. This study provides a rapid, simple, and engaging method to attribute the burden of animal health loss at a high-level. Results are informative, however will become increasingly useful once they can be compared with results from more sophisticated, data-driven models.

Competition and sex-age class alter the effects of group size on vigilance in white-tailed deer Odocoileus virginianus

Competition and sex-age class alter the effects of group size on vigilance in white-tailed deer Odocoileus virginianus

Examining postmounting grooming in male golden snub-nosed monkeys to investigate the functions of same-sex mounts.

Same-sex mounting behavior has been documented across primate species and several biological functions, that are often difficult to differentiate, have been proposed. Postmounting affiliative behaviors (e.g., embracing and grooming) have been partly overlooked and their performance may more clearly reveal the function of same-sex mounts for different age-sex classes. Here, considering postmounting grooming behaviors (PMG), we tested the affiliation hypothesis for same-sex mounts in an all-male unit (AMU) of captive golden snub-nosed monkeys (Rhinopithecus roxellana) (n = 7-8) and the needing to learn hypothesis in the juveniles of the AMU (n = 4) and the juvenile male from the neighboring one-male unit in Shanghai Wild Animal Park, China from November 2014 to June 2015. A total of 1986 same-sex mounts were recorded from the AMU individuals and the juvenile of the neighboring one-male unit. We found that neither dyadic proximity-based association nor grooming-based affiliation predicted the occurrence of mounts in the AMU, and PMG was more likely to occur in nontense than tense social contexts, providing no support for the affiliation hypothesis. Although dyadic grooming-based affiliation predicted the occurrence of PMG, it exerted no influence on the occurrence of mounts, which was necessary to support the hypothesis. However, consistent with the needing to learn hypothesis, from ages two to four, juveniles' mounting duration increased and they performed pelvic thrusting during mounting more and more often as they grew older. Reciprocated series mounts were much more common among juveniles than other mounting dyads, providing learning opportunities for both participants. The mounter was more likely to groom the mountee at the end of a mount among juveniles than other mounting dyads, suggesting that juvenile mounters may profit from mounting and groom mountees in return for providing them with opportunities to learn copulatory skills. A hypothesis that merits further investigation.

Population demography of the Blackbuck Antilope cervicapra (Cetartiodactyla: Bovidae) at Point Calimere Wildlife Sanctuary, India

Demographic research utilizing vital rates and life tables is a standard aspect of planning protection and management strategies for wildlife populations. The Blackbuck Antilope cervicapra population at Point Calimere Wildlife Sanctuary, Tamil Nadu, has undergone fluctuations in recent decades. The cause remains elusive, and conservation efforts may be hampered by a lack of population data. This study aimed to estimate demographic parameters using population count and age-sex classification data collected for the years 2017─2020. The overall mean population estimate derived from line-transect distance sampling was 719, with annual estimates of 716, 727, 711, and 722 for the years 2017–2020 respectively. In total, 64% of Blackbucks counted were adults, 19% subadults, and 17% fawns. Mortality was highest for adult and subadult classes for the composite female class, and fawn mortality was 20%. The net reproductive rate (R0) was as low as 3.28 offspring per generation contrasted with a rather longer mean generation time (G) as 4.75 years. Thus, the study observes a decrease in Blackbuck numbers postulated in earlier research to be driven principally by a conglomerate of factors, including reduction of usable space and interspecific resource competition. Our findings provide a baseline demography of the species and highlights the value of long-term demographic monitoring of age sex classes to understand the evolution of life histories.

Reciprocity and beyond: Explaining meat transfers in savanna-dwelling chimpanzees at Fongoli, Senegal.

To understand the function of food sharing among our early hominin ancestors, we can turn to our nonhuman primate relatives for insight. Here, we examined the function of meat sharing by Fongoli chimpanzees, a community of western chimpanzees (Pan troglodytes verus) in southeastern Sénégal. We tested three non-mutually exclusive hypotheses that have been used to explain patterns of food sharing: kin selection, generalized reciprocity, and meat-for-mating opportunities. We analyzed meat sharing events (n = 484) resulting from hunts, along with data on copulations, age-sex class, and kinship to determine which variables predict the likelihood of meat sharing during this study period (2006-2019). We found full or partial support for kin selection, direct reciprocity, and meat-for-mating-opportunities. However, the analyses reveal that reciprocity and a mother/offspring relationship were the strongest predictors of whether or not an individual shared meat. The results of this study emphasize the complexity of chimpanzee meat sharing behaviors, especially at a site where social tolerance offers increased opportunities for meat sharing by individuals other than dominant males. These findings can be placed in a referential model to inform hypotheses about the sensitivity of food sharing to environmental pressures, such as resource scarcity in savanna landscapes.

Age differences in the responses of vervet monkeys, Chlorocebus pygerythrus, to terrestrial alarm calls

The high costs of predation and the opportunity costs associated with predator avoidance are likely to select for flexibility in the development of antipredator responses based on local socioecological conditions. As group size is hypothesized to vary across populations as a function of predation risk, the development of antipredator behaviours throughout ontogeny may be influenced by the size of an individual's social group. Here, we explore the development of alarm call responses in wild vervet monkeys living at the Samara Game Reserve, South Africa. Vervet monkeys at this site live in relatively large social groups, and adult responses to alarms have previously shown less uniformity than in other sites where group sizes are smaller. We presented monkeys playback recordings of terrestrial alarm calls produced by individuals of different age–sex classes. We then videorecorded and scored the responses of receivers along an ordinal maturity scale. We used a mixed-effects ordinal logistic regression model within a Bayesian framework to explore how response intensity is affected by the age–sex of the caller, and the age of receivers. Our analysis showed that younger monkeys (<2 years old) exhibit strong evasive responses to call stimuli, regardless of the age–sex class of the caller. The intensity of these responses decreases with age, with responses to nonalarm calls decreasing earlier in development than responses to alarm calls. Adult responses to alarm calls in this population are as likely to be characterized by a general increase in vigilance as they are to consist of an evasive response. We suggest that responses in younger individuals at Samara are mediated by generalized startle responses to loud, plosive noises. Development of more specific responses throughout ontogeny is likely to be dependent on learning from adult models, whose milder responses reflect local socioecological conditions specific to Samara.

Why hibernate? Tests of four hypotheses to explain intraspecific variation in hibernation phenology

Abstract Hibernation is a remarkable behaviour deployed by a diverse array of endotherms within many clades that greatly reduces metabolic need, but also has somatic costs. Hibernation in modern endotherms is often assumed to be an adaptation allowing animals to avoid extreme thermal conditions or food shortages in seasonal environments. However, many animals hibernate when foraging conditions are energetically profitable, suggesting other causal factors influence hibernation behaviour. Understanding the selection pressures responsible for intraspecific variation in the timing and duration of hibernation can help elucidate the relative evolutionary influences of the ultimate ecological causes of hibernation. We tested four previously proposed mechanistic hypotheses to explain intraspecific variation in hibernation phenology in the federally threatened northern Idaho ground squirrel (Urocitellus brunneus): (1) thermal tolerance, (2) food limitation, (3) predation avoidance and (4) sexual selection. The predation avoidance and sexual selection hypotheses received the most support, although we also found some support for the thermal tolerance and food limitation hypotheses. Heavy squirrels increased hibernation duration regardless of environmental conditions, as predicted solely by the predation avoidance hypothesis. Reproductive males emerged from hibernation earlier in spring than other sex–age classes, a pattern predicted by the sexual selection hypothesis. Temperature and food availability explained a much smaller amount of the variation in hibernation behaviour, only partially supporting predictions of the thermal tolerance and food limitation hypotheses. Our results indicate that animals navigate life‐history trade‐offs between energetic allocation to survival and reproduction via state‐dependent optimization of hibernation phenology. Consequently, any future environmental changes that influence body condition will have implications for population ecology and life‐history evolution of hibernating animals due to stark differences in daily survival probability between hibernation and the active season. Read the free Plain Language Summary for this article on the Journal blog.

Quantifying the movement, behaviour and environmental context of group-living animals using drones and computer vision.

Methods for collecting animal behaviour data in natural environments, such as direct observation and biologging, are typically limited in spatiotemporal resolution, the number of animals that can be observed and information about animals' social and physical environments. Video imagery can capture rich information about animals and their environments, but image-based approaches are often impractical due to the challenges of processing large and complex multi-image datasets and transforming resulting data, such as animals' locations, into geographical coordinates. We demonstrate a new system for studying behaviour in the wild that uses drone-recorded videos and computer vision approaches to automatically track the location and body posture of free-roaming animals in georeferenced coordinates with high spatiotemporal resolution embedded in contemporaneous 3D landscape models of the surrounding area. We provide two worked examples in which we apply this approach to videos of gelada monkeys and multiple species of group-living African ungulates. We demonstrate how to track multiple animals simultaneously, classify individuals by species and age-sex class, estimate individuals' body postures (poses) and extract environmental features, including topography of the landscape and animal trails. By quantifying animal movement and posture while reconstructing a detailed 3D model of the landscape, our approach opens the door to studying the sensory ecology and decision-making of animals within their natural physical and social environments.

Behavioural responses of humpback whales (Megaptera novaeangliae) to whalewatching boats near Isla de la Plata, Machalilla National Park, Ecuador

Machalilla National Park, on the coast of mainland Ecuador, supports a growing whalewatching industry that focuses on Southern Hemisphere humpback whales, which spend the austral winter (June-September) in this area. This study was designed to measure short-term reactions of whales to the whalewatching vessel activity typically seen in this area for two reasons: (1) to identify the nature of whales’ avoidance response, if any, in order to draft whalewatching guidelines that help local mariners identify when they may be disturbing whales; and (2) to quantify the magnitude of any avoidance response, to examine how this relatively understudied population behaves around boats compared with whales in other whalewatching areas. A shore-based theodolite tracking team created a ‘natural’ experiment to observe relationships between whalewatching traffic and whale behaviour in 1998 and 1999. Swim speed and path directness of humpback whales were measured in the absence of boats, and how those parameters changed when boats arrived was recorded. When whales entered the study area accompanied by boats, a record was made of how their behaviour changed after the boats left. Humpback whales reacted to the approach of whalewatching boats by increasing swim speed significantly, and adopted a much more direct path after boats left. Future research is needed to determine whether responses vary with number, proximity or type of vessel. Similarly, future studies are recommended to determine whether different age-sex classes vary in vulnerability to disturbance. Meanwhile, this study enables provision of much-needed, practical advice to local operators who are concerned that they may be disturbing whales: one way that mariners can tell if they are causing disturbance is if they need to increase their vessel’s speed to keep pace. The average behavioural responses measured were strong enough to recommend that Machalilla National Park adopt precautionary management procedures to limit number and proximity of vessels.

Non-Reproductive Sexual Behavior in Wild White-Thighed Colobus Monkeys (Colobus vellerosus).

Rare behaviors are often missing from published papers, hampering phylogenetic analyses. Here, we report, for the first time, masturbation and same-sex sexual behavior (SSB) in both male and female black-and-white colobus monkeys. We recorded these behaviors during 32months of observation (1573h of focal animal sampling) on Colobus vellerosus collected at the Boabeng-Fiema Monkey Sanctuary in Ghana. Males were observed masturbating and involved in SSB more than females. Subadult males were the age-sex class that engaged in both of these behaviors most often and a third of all SSB observed in young males occurred when they were forming an all-male band (AMB), which are temporally transient social groups in this species. Our data support that masturbation in males may be a sexual outlet for individuals that do not have a current sexual partner, while in females it may function in mate attraction by advertising receptivity. SSB may occur as an evolutionary byproduct but given the temporal clustering of observed events in males prior to AMB formation, our data best support the hypothesis that these behaviors facilitate male-male bonding (i.e., act as social glue). Within AMB's, males engage in coalitionary behavior to take over social groups containing females and strong bonds are important for success and later access to females, which could have selected for SSB in C. vellerosus.

Brown bear skin-borne secretions display evidence of individuality and age-sex variation

Scent originates from excretions and secretions, and its chemical complexity in mammals translates into a diverse mode of signalling. Identifying how information is encoded can help to establish the mechanisms of olfactory communication and the use of odours as chemical signals. Building upon existing behavioural and histological literature, we examined the chemical profile of secretions used for scent marking by a solitary, non-territorial carnivore, the brown bear (Ursus arctos). We investigated the incidence, abundance, and uniqueness of volatile organic compounds (VOCs) from cutaneous glandular secretions of 12 wild brown bears collected during late and post-breeding season, and assessed whether age-sex class, body site, and individual identity explained profile variation. VOC profiles varied in the average number of compounds, compound incidence, and compound abundance by age-sex class and individual identity (when individuals were grouped by sex), but not by body site. Mature males differed from other age-sex classes, secreting fewer compounds on average with the least variance between individuals. Compound uniqueness varied by body site and age for both males and females and across individuals. Our results indicate that brown bear skin-borne secretions may facilitate age-sex class and individual recognition, which can contribute towards further understanding of mating systems and social behaviour.