Expectations versus Reality of Designer Dog Ownership in the United States.

"Designer dogs", which are the hybrid offspring that result from intentionally breeding dogs belonging to different breeds, are an extremely popular pet choice in the United States. Poodle mixes, often called "doodles", are a very common type of designer dog. However, there are many misconceptions surrounding them, and the reality of owning one may not match the owner's expectations. For instance, many people believe these dogs to be non-shedding and hypoallergenic, although this is not always the case. This study explored whether the reality of owning a doodle matches owner expectations. For comparison purposes, we also asked owners of non-doodle dogs about their expectations versus reality. Our survey-based study included 2191 owners of doodles and non-doodle dogs recruited via groups of dog owners on Facebook and Reddit. The data showed that, when selecting their dogs, doodle owners were more influenced than non-doodle owners by their dog's appearance and by the perception that doodles are good with children and are generally healthy. Doodle owners reported being highly satisfied with their dogs; nevertheless, more than twice as many doodle owners than owners of the other groups of dogs reported that their dog's maintenance requirements, such as their need for regular grooming, were more intensive than they had expected. This finding suggests that those interested in owning doodles would benefit from having more information about their dog's grooming needs so they can decide whether they have the time and money required to meet their dog's welfare needs.

Characterizing Pet Acquisition and Retention During the COVID-19 Pandemic.

In March 2020, Americans began experiencing numerous lifestyle changes due to the COVID-19 pandemic. Some reports have suggested that pet acquisition and ownership increased during this period, and some have suggested shelters and rescues will be overwhelmed once pandemic-related restrictions are lifted and lifestyles shift yet again. In May 2021, the ASPCA hired the global market research company Ipsos to conduct a general population survey that would provide a more comprehensive picture of pet ownership and acquisition during the pandemic. Although pet owners care for a number of species, the term pet owner in this study specifically refers to those who had dogs and/or cats. One goal of the survey was to determine whether data from a sample of adults residing in the United States would corroborate findings from national shelter databases indicating that animals were not being surrendered to shelters in large numbers. Furthermore, this survey gauged individuals' concerns related to the lifting of COVID-19 restrictions, and analyses examined factors associated with pet owners indicating they were considering rehoming an animal within the next 3 months. The data showed that pet ownership did not increase during the pandemic and that pets may have been rehomed in greater numbers than occurs during more stable times. Importantly, rehomed animals were placed with friends, family members, and neighbors more frequently than they were relinquished to animal shelters and rescues. Findings associated with those who rehomed an animal during the pandemic, or were considering rehoming, suggest that animal welfare organizations have opportunities to increase pet retention by providing resources regarding pet-friendly housing and affordable veterinary options and by helping pet owners strategize how to incorporate their animals into their post-pandemic lifestyles.

The Experience of Teleworking with Dogs and Cats in the United States during COVID-19.

In Spring of 2020, the novel coronavirus (SAR-CoV-2) prompted an unprecedented number of individuals across the United States to begin working from home. Prior research has identified both positive and negative impacts of teleworking on employee well-being, and this study built on that research to explore perceptions regarding how companion animals factor into the teleworking experience. Individuals who had experience working from home and from their employer's office completed an online survey about those experiences. Participants reported spending more quality time with their companion animals and family members when they worked from home. Furthermore, when working from home, individuals with dogs were more likely than those without dogs to report they socialized with other people, got a healthy amount of physical activity, and took at least one 15-min walk during the workday. Some participants, particularly those in households containing both dogs and cats, indicated that their pets created distractions during the workday. Future studies can build on this research by investigating whether the findings persist once the novel coronavirus is no longer a threat, and by paying close attention to the characteristics of pets, owners, and household dynamics that may influence the effects of pet ownership on the teleworking experience.

Assessing the Impact of a Virtual Shelter Medicine Rotation on Veterinary Students' Knowledge, Skills, and Attitudes Regarding Access to Veterinary Care.

Strong bonds commonly form between companion animals and people of all socio-demographic backgrounds, yet many pet owners face numerous barriers to accessing veterinary care for their companion animals. For example, they may have difficulties paying for care; they may lack veterinary practices in their community; and they may experience language barriers that impede their ability to utilize veterinary services. Various strategies exist that can help veterinarians address the diverse needs of pet owners in their communities, but these techniques are not commonly covered in the veterinary school curriculum. This study explored how including in-depth, purposefully curated information about access to veterinary care issues within a required shelter medicine rotation impacted fourth-year veterinary students' knowledge, skills, and attitudes regarding the problems clients commonly face when seeking access to veterinary care. Students participated either in a control group of a virtual, four-week rotation delivered via Zoom meetings and self-study, or in an experimental group that additionally completed an interactive online learning module. The online module heavily featured issues surrounding access to veterinary care. Irrespective of which version of the rotation students enrolled, their opinions grew more favorable from pretest to post-test regarding the role of not-for-profit veterinary clinics in communities, as did their expectations that veterinarians should provide affordable treatment options. Additionally, students in the experimental group demonstrated from pretest to post-test increased awareness of the potential for implicit bias toward pet owners within veterinary practice and showed a reduction in their tendency to be judgmental of veterinary clients. By the end of the study, students in the experimental group also expressed greater confidence in their ability to offer incremental care treatment options to veterinary clients. These findings suggest that providing content that focuses on increasing access to veterinary care enhances students' awareness of the need to offer a variety of treatment and payment options to clients. Findings from this study can inform curriculum design in veterinary schools and continuing education programs for veterinary professionals.

Human-Animal Co-Sleeping: An Actigraphy-Based Assessment of Dogs' Impacts on Women's Nighttime Movements.

Humans regularly enter into co-sleeping arrangements with human and non-human partners. Studies of adults who co-sleep report that co-sleeping can impact sleep quality, particularly for women. Although dog owners often choose to bedshare with their dogs, we know relatively little about the nature of these relationships, nor the extent to which co-sleeping might interfere with sleep quality or quantity. In an effort to rectify this, we selected a sample of 12 adult female human (M = 50.8 years) and dog dyads, and monitored their activity using actigraphy. We collected movement data in one-minute epochs for each sleep period for an average of 10 nights per participant. This resulted in 124 nights of data, covering 54,533 observations (M = 7.3 hours per night). In addition, we collected subjective sleep diary data from human participants. We found a significant positive relationship between human and dog movement over sleep periods, with dogs influencing human movement more than humans influenced dog movement. Dog movement accompanied approximately 50% of human movement observations, and dog movement tripled the likelihood of the human transitioning from a non-moving state to a moving state. Nevertheless, humans rarely reported that their dog disrupted their sleep. We encourage the continued exploration of human-animal co-sleeping in all its facets and provide recommendations for future research in this area.

A Combined Approach to Predicting Rest in Dogs Using Accelerometers.

The ability to objectively measure episodes of rest has clear application for assessing health and well-being. Accelerometers afford a sensitive platform for doing so and have demonstrated their use in many human-based trials and interventions. Current state of the art methods for predicting sleep from accelerometer signals are either based on posture or low movement. While both have proven to be sensitive in humans, the methods do not directly transfer well to dogs, possibly because dogs are commonly alert but physically inactive when recumbent. In this paper, we combine a previously validated low-movement algorithm developed for humans and a posture-based algorithm developed for dogs. The hybrid approach was tested on 12 healthy dogs of varying breeds and sizes in their homes. The approach predicted state of rest with a mean accuracy of 0.86 (SD = 0.08). Furthermore, when a dog was in a resting state, the method was able to distinguish between head up and head down posture with a mean accuracy of 0.90 (SD = 0.08). This approach can be applied in a variety of contexts to assess how factors, such as changes in housing conditions or medication, may influence a dog's resting patterns.

Dog rivalry impacts following behavior in a decision-making task involving food.

Dogs learn a great deal from humans and other dogs. Previous studies of socially influenced learning between dogs have typically used a highly trained demonstrator dog who is unfamiliar to the observer. Because of this, it is unknown how dynamics between familiar dogs may influence their likelihood of learning from each other. In this study, we tested dogs living together in two-dog households on whether individual dogs' rivalry scores were associated with performance on a local enhancement task. Specifically, we wanted to know whether dog rivalry impacted whether an observer dog would approach a plate from which a demonstrator dog had eaten all available food, or whether the observer dog would approach the adjacent plate that still contained food. Dog rivalry scores were calculated using the Canine Behavioral Assessment and Research Questionnaire and indicated each dog's tendency to engage aggressively with the other household dog. Low-rivalry dogs were more likely to approach the empty plate than high-rivalry dogs when the observer dog was allowed to approach the plates immediately after the demonstrator had moved out of sight. This difference between low- and high-rivalry dogs disappeared, however, when observer dogs had to wait 5 s before approaching the plates. The same pattern was observed during a control condition when a human removed the food from a plate. Compared to low-rivalry dogs, high-rivalry dogs may pay less attention to other dogs due to a low tolerance for having other dogs in close proximity.

Dogs on the Move: Factors Impacting Animal Shelter and Rescue Organizations' Decisions to Accept Dogs from Distant Locations.

Long-distance dog transfer programs are a topic of burgeoning interest in the animal welfare community, but little research has focused on such programs. This exploratory study, which surveyed 193 individuals associated with animal shelter and rescue organizations in the United States, evaluated factors that impacted organizations' decisions to transfer in dogs over long distances (>100 miles) and assessed what criteria were commonly valued by destination organizations. Specifically, we examined the following aspects of long-distance transfer programs: (1) logistics of long-distance dog transfers; (2) factors impacting dog selection; (3) medical requirements; (4) partnerships formed between source and destination organizations; and (5) perceptions of long-distance dog transfer programs by individuals affiliated with the destination organizations. This study revealed that many logistical considerations factor into transfer decisions and the formation of healthy partnerships between source and destination organizations. Participants indicated their organization's willingness to receive dogs of various sizes, coat colors and ages, but organizations often had restrictions regarding the breeds they would accept. Study findings indicate some organizations have strict quarantine policies and pre-transfer medical requirements, while others have no such requirements.

An Evaluation of the Role the Internet Site Petfinder Plays in Cat Adoptions.

To better understand factors contributing to cat adoptions, this study was used to explore a possible association between an adoptable cat's popularity on Petfinder and the cat's length of availability for adoption at a managed-intake nonhuman animal shelter. This study was also used to examine factors associated with a cat's popularity on Petfinder and the percentage of adopters who visited Petfinder before making adoption decisions. One third of adopters surveyed visited Petfinder before adopting, and half of those had viewed their adopted cats' Petfinder profiles. The number of clicks per day that cats received on the site was negatively correlated with their length of availability. Age at adoption was positively correlated with length of availability and negatively correlated with number of clicks per day. Coat color was a strong predictor of number of clicks per day and length of availability. The only variable within the photographer's control significantly associated with number of clicks per day was whether the photos included toys. Although cats' physical characteristics are strong predictors of their popularity, strategic use of toys in cats' photographs may promote adoptions of cats who are typically overlooked.

Is that dog a pit bull? A cross-country comparison of perceptions of shelter workers regarding breed identification.

Bull breeds are commonly kept as companion animals, but the pit bull terrier is restricted by breed-specific legislation (BSL) in parts of the United States and throughout the United Kingdom. Shelter workers must decide which breed(s) a dog is. This decision may influence the dog's fate, particularly in places with BSL. In this study, shelter workers in the United States and United Kingdom were shown pictures of 20 dogs and were asked what breed each dog was, how they determined each dog's breed, whether each dog was a pit bull, and what they expected the fate of each dog to be. There was much variation in responses both between and within the United States and United Kingdom. UK participants frequently labeled dogs commonly considered by U.S. participants to be pit bulls as Staffordshire bull terriers. UK participants were more likely to say their shelters would euthanize dogs deemed to be pit bulls. Most participants noted using dogs' physical features to determine breed, and 41% affected by BSL indicated they would knowingly mislabel a dog of a restricted breed, presumably to increase the dog's adoption chances.

Do Dog Behavioral Characteristics Predict the Quality of the Relationship between Dogs and Their Owners?

This paper explores whether dog behavioral characteristics predict the quality of the relationship between dogs and their owners (i.e., owner attachment to dog), and whether relations between dog behavior and owner attachment are moderated by demographic characteristics. In this study, N = 92 children and N = 60 adults from 60 dog-owning families completed questionnaires about their attachment to their pet dog, their level of responsibility for that dog, and their general attitudes toward pets. They also rated their dogs on observable behavioral characteristics. Individuals who held positive attitudes about pets and who provided much of their dog's care reported stronger attachments to their dogs. The strength of owners' attachments to their dogs was associated with dog trainability and separation problems. Relationships between owner attachment and both dog excitability and attention-seeking behavior were further moderated by demographic characteristics: for Caucasians but not for non-Caucasians, dog excitability was negatively associated with owner attachment to dog; and for adults, dog attention-seeking behavior was positively associated with owner attachment, but children tended to be highly attached to their dogs, regardless of their dogs' attention-seeking behaviors. This study demonstrates that certain dog behavioral traits are indeed associated with the strength of owners' attachments to their dogs.

Genetic and Environmental Influences on Individual Differences in Frequency of Play with Pets among Middle-Aged Men: A Behavioral Genetic Analysis.

There is growing evidence that pet ownership and human-animal interaction (HAI) have benefits for human physical and psychological well-being. However, there may be pre-existing characteristics related to patterns of pet ownership and interactions with pets that could potentially bias results of research on HAI. The present study uses a behavioral genetic design to estimate the degree to which genetic and environmental factors contribute to individual differences in frequency of play with pets among adult men. Participants were from the ongoing longitudinal Vietnam Era Twin Study of Aging (VETSA), a population-based sample of 1,237 monozygotic (MZ) and dizygotic (DZ) twins aged 51-60 years. Results demonstrate that MZ twins have higher correlations than DZ twins on frequency of pet play, suggesting that genetic factors play a role in individual differences in interactions with pets. Structural equation modeling revealed that, according to the best model, genetic factors accounted for as much as 37% of the variance in pet play, although the majority of variance (63-71%) was due to environmental factors that are unique to each twin. Shared environmental factors, which would include childhood exposure to pets, overall accounted for <10% of the variance in adult frequency of pet play, and were not statistically significant. These results suggest that the effects of childhood exposure to pets on pet ownership and interaction patterns in adulthood may be mediated primarily by genetically-influenced characteristics.

Chronic stress, allostatic load, and aging in nonhuman primates.

Allostatic load is the "wear and tear" of the body resulting from the repeated activation of compensatory physiological mechanisms in response to chronic stress. Allostatic load can significantly affect the aging process and result in reduced longevity, accelerated aging, and impaired health. Although low socioeconomic status is associated with high allostatic load during aging, the effects of status-related psychosocial stress on allostatic load are often confounded by lifestyle variables. Chronic psychosocial stress associated with low dominance rank in nonhuman primates represents an excellent animal model with which to investigate allostatic load and aging in humans. Research conducted with free-ranging rhesus monkeys suggests that female reproduction can also be a source of stress and allostatic load. Female reproduction is associated with increased risk of mortality and hyperactivation of the hypothalamic-pituitary-adrenal axis. Reproduction is especially stressful and costly for aging females of low rank. Although many indicators of body condition and neuroendocrine and immune function are influenced by aging, there are marked and stable individual differences among aging females in body condition, plasma cortisol responses to stress, and cytokine responses to stress. These differences are consistent with the hypothesis that there are strong differences in chronic stress among individuals, and that allostatic load resulting from chronic stress affects health during aging. Comparisons between captive and free-ranging rhesus monkey populations may allow us to understand how differences in environmental stress and allostatic load affect rates of aging, and how these in turn translate into differences in longevity and health.

Immune function and HPA axis activity in free-ranging rhesus macaques.

In mammals, the hypothalamic-pituitary-adrenal (HPA) axis and immune system play an important role in the maintenance of homeostasis. Dysregulation of either system resulting, for example, from psychosocial or reproductive stress increases susceptibility to disease and mortality risk, especially in aging individuals. In a study of free-ranging rhesus macaques, we examined how female age, reproductive state, social rank, and body condition influence (i) aspects of cytokine biology (plasma concentrations of interleukin-1 receptor antagonist (IL-1ra), IL-6 and IL-8), and (ii) HPA axis activity (plasma and fecal glucocorticoid levels). We also assessed individual differences in cytokine and hormone concentrations over time to determine their consistency and to investigate relations between these two indicators of physiological regulation and demand. Female monkeys showed marked increases in HPA axis activity during pregnancy and lactation, and increased circulating levels of IL-1ra with advancing age. Inter-individual differences in IL-1ra and IL-8 were consistent over successive years, suggesting that both are stable, trait-like characteristics. Furthermore, the concentrations of fecal glucocorticoid hormones in non-pregnant, non-lactating females were correlated with their plasma cortisol and IL-8 concentrations. Some individuals showed permanently elevated cytokine levels or HPA axis activity, or a combination of the two, suggesting chronic stress or disease. Our results enhance our understanding of within- and between-individual variation in cytokine levels and their relationship with glucocorticoid hormones in free-ranging primates. These findings can provide the basis for future research on stress and allostatic load in primates.

Mu-opioid receptor (OPRM1) variation, oxytocin levels and maternal attachment in free-ranging rhesus macaques Macaca mulatta.

Understanding the genetic and neuroendocrine basis of the mother-infant bond is critical to understanding mammalian affiliation and attachment. Functionally similar nonsynonymous mu-opioid receptor (OPRM1) SNPs have arisen and been maintained in humans (A118G) and rhesus macaques Macaca mulatta (C77G). In rhesus macaques, variation in OPRM1 predicts individual differences in infant affiliation for mothers. Specifically, infants carrying the G allele show increased distress on separation from their mothers, and spend more time with them upon reunion, than individuals homozygous for the C allele. In humans, individuals possessing the G allele report higher perceptions of emotional pain on receiving rejection by social partners. We studied maternal behavior over the course of a year among free-ranging female rhesus macaques on Cayo Santiago, Puerto Rico. We then trapped females and collected blood samples from which we assessed OPRM1 genotype; we also collected cerebrospinal fluid samples from which we measured oxytocin (OT) levels. We show that females possessing the G allele restrain their infants more (i.e., prevent infants from separating from them by pulling them back) than females homozygous for the C allele. Females possessing the G allele also show higher OT levels when lactating, and lower OT levels when neither lactating nor pregnant, than females homozygous for the C allele. This is the first study to demonstrate an association between OPRM1 genotype and maternal attachment for infants, and is one of the first studies of any free-ranging primate population to link functional genetic variation to behavior via potentially related neuroendocrine mechanisms.

Effects of age on cerebrospinal fluid oxytocin levels in free-ranging adult female and infant rhesus macaques.

There is growing interest in examining oxytocin and social functioning in human and non-human primates. Studies of human oxytocin biology are typically restricted to peripheral assessments because opportunities to collect cerebrospinal fluid (CSF) are rare. Several studies have examined CSF oxytocin levels in captive adult primates, but none to our knowledge have been conducted under free-ranging conditions and inclusive of infants. The main goal of this study was to establish feasibility of quantifying CSF oxytocin levels in free-ranging adult female and infant rhesus monkeys living on Cayo Santiago, PR. CSF oxytocin levels were examined in relation to individuals' demographic and reproductive characteristics as well as plasma cortisol levels. CSF oxytocin concentrations ranged from 36.02 to 134.41 pg/ml in adult females (ages 7-26 years; N = 31) and 35.94 to 77.3 pg/ml in infants (ages 38-134 days; N = 17). CSF oxytocin levels were positively correlated with adult female age and negatively correlated with infant age. The former correlation was driven by reproductive status. CSF oxytocin levels were unrelated to dominance rank or plasma cortisol levels. In contrast to a previous study of plasma oxytocin concentrations in this population, CSF oxytocin levels did not differ significantly between lactating and non-lactating females. These findings: 1) provide feasibility data for examining CSF oxytocin levels in free-ranging non-human primates and 2) indicate that CSF oxytocin levels may be a biomarker of age-related central nervous system changes across lifespan development. Research is now required to examine CSF oxytocin levels in the context of social functioning in free-ranging rhesus monkeys.

Terminal investment and senescence in rhesus macaques (Macaca mulatta) on Cayo Santiago.

Long-lived iteroparous species often show aging-related changes in reproduction that may be explained by 2 non-mutually exclusive hypotheses. The terminal investment hypothesis predicts increased female reproductive effort toward the end of the life span, as individuals have little to gain by reserving effort for the future. The senescence hypothesis predicts decreased female reproductive output toward the end of the life span due to an age-related decline in body condition. Nonhuman primates are ideal organisms for testing these hypotheses, as they are long lived and produce altricial offspring heavily dependent on maternal investment. In this study, we integrated 50 years of continuous demographic records for the Cayo Santiago rhesus macaque (Macaca mulatta) population with new morphometric and behavioral data to test the senescence and terminal investment hypotheses. We examined relationships between maternal age and activity, mother and infant body condition, interbirth intervals, measures of behavioral investment in offspring, and offspring survival and fitness to test for age-associated declines in reproduction that would indicate senescence, and for age-associated increases in maternal effort that would indicate terminal investment. Compared with younger mothers, older mothers had lower body mass indices and were less active, had longer interbirth intervals, and spent more time in contact with infants, but had infants of lower masses and survival rates. Taken together, our results provide strong evidence for the occurrence of reproductive senescence in free-ranging female rhesus macaques but are also consistent with some of the predictions of the terminal investment hypothesis.

Effects of reproductive condition and dominance rank on cortisol responsiveness to stress in free-ranging female rhesus macaques.

The hypothalamic-pituitary-adrenal (HPA) axis modulates individuals' physiological responses to social stress, which is an inevitable aspect of the daily lives of group-living animals. Previous nonhuman primate studies have reported that sex, age, rank, and reproductive condition influence cortisol levels under stressful conditions. In this study we investigated cortisol responses to stress among 70 multiparous, free-ranging female rhesus macaques (Macaca mulatta) on the island of Cayo Santiago, PR. Plasma cortisol samples were collected in two consecutive years under similar conditions. Twenty-two females were sampled both years, and most of those females were lactating in only one of the years. Individual differences in cortisol levels were stable across years, even though reproductive condition changed for most individuals. No relationship was found between age or social rank and cortisol levels. Of the females that changed reproductive conditions, cortisol levels were higher when they were lactating than when they were cycling, and the amount of change in cortisol from cycling to lactating was greatest for low-ranking individuals. Heightened reactivity to stress during lactation may be the result of concerns about infant safety, and such concerns may be higher among low-ranking mothers than among higher ranking mothers. Psychosocial stress and hyperactivation of the HPA axis during lactation can suppress immune function and increase vulnerability to infectious diseases, thus explaining why adult females in the free-ranging rhesus macaque population on Cayo Santiago have a higher probability of mortality during the birth season than during the mating season.

Mother-infant interactions in free-ranging rhesus macaques: relationships between physiological and behavioral variables.

Studies of mother-infant relationships in nonhuman primates have increasingly attempted to understand the neuroendocrine bases of interindividual variation in mothering styles and the mechanisms through which early exposure to variable mothering styles affects infant behavioral development. In this study of free-ranging rhesus macaques on Cayo Santiago, Puerto Rico, we aimed to: 1) compare lactating and nonlactating females to investigate whether lactation is associated with changes in plasma cortisol, prolactin and oxytocin, as well as changes in CSF levels of serotonin and dopamine metabolites (5-HIAA and HVA); 2) examine the extent to which interindividual variation in maternal physiology is associated with variation in maternal behavior; 3) examine the extent to which interindividual variation in infant physiology and behavior is accounted for by variation in maternal physiology and behavior. Lactating females had higher plasma concentrations of cortisol, prolactin, and oxytocin but lower CSF concentrations of HVA than nonlactating females. Variation in maternal rejection behavior was positively correlated with variation in maternal plasma cortisol levels and in CSF 5-HIAA levels while variation in the time spent nursing and grooming was associated with maternal plasma oxytocin levels. Infants who were protected more by their mothers had higher cortisol levels than those who were protected less, while infants who were rejected more had lower CSF 5-HIAA than infants who were rejected less. Since exposure to high levels of maternal protectiveness and rejection is known to affect the offspring's behavior and responsiveness to the environment later in life, our results are consistent with the hypothesis that these effects are mediated by long-term changes in the activity of the offspring's HPA axis and brain serotonergic system.

Sex differences in survival costs of reproduction in a promiscuous primate.

In sexually promiscuous mammals, female reproductive effort is mainly expressed through gestation, lactation, and maternal care, whereas male reproductive effort is mainly manifested as mating effort. In this study, we investigated whether reproduction has significant survival costs for a seasonally breeding, sexually promiscuous species, the rhesus macaque, and whether these costs occur at different times of the year for females and males, namely in the birth and the mating season, respectively. The study was conducted with the rhesus macaque population on Cayo Santiago, Puerto Rico. Data on 7,402 births and 922 deaths over a 45-year period were analyzed. Births were concentrated between November and April, while conceptions occurred between May and October. As predicted, female mortality probability peaked in the birth season whereas male mortality probability peaked in the mating season. Furthermore, as the onset of the birth season gradually shifted over the years in relation to climatic changes, there was a concomitant shift in the seasonal peaks of male and female mortality. Taken together, our findings provide the first evidence of sex differences in the survival costs of reproduction in nonhuman primates and suggest that reproduction has significant fitness costs even in environments with abundant food and absence of predation.