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Objectives: Existing data on bonobo and chimpanzee dental eruption timing are derived predominantly from captive individuals or deceased wild individuals. However, recent advances in noninvasive photographic monitoring of living, wild apes have enabled researchers to characterize dental eruption in relatively healthy individuals under naturalistic conditions. At present, such data are available for only one population of wild chimpanzees. We report data for an additional population of wild chimpanzees and the first dental eruption data for wild bonobos. Materials and Methods: We collected photographs and video footage of teeth from the open mouths of wild bonobos and East African chimpanzees of known age from LuiKotale, Democratic Republic of the Congo, and Gombe National Park, Tanzania, respectively. We scored the presence and absence of deciduous teeth from photographs and video footage to characterize deciduous dental eruption timing in these two populations. Results: Deciduous dental eruption ages in our sample fall within the range of variation previously documented for captive chimpanzees, but eruption ages are later in wild than in captive contexts. We found substantial variation in deciduous canine eruption timing, particularly among bonobos. One bonobo had a deciduous canine present by 227 days old while another did not have a deciduous canine present at 477 days old. Discussion Our data indicate that deciduous teeth erupt later in wild individuals than in captive individuals. We also found that deciduous dental eruption timing varies considerably between individuals within our study populations, a pattern that is consistent with previous studies. Future studies should consider sources of variation in deciduous canine eruption timing and relationships with other aspects of life history as additional data become available. [ABSTRACT FROM AUTHOR]

Highly differentiated and affiliative social relationships are observed in a variety of mammals, including primates, cetaceans, and social carnivores. Although there has been a transformation in our understanding of the form and function of social bonds in the past two decades, the role of early life social experiences in the development of bonds remains less clear. Few studies have examined whether social relationships during infancy and juvenility (aside from those between mothers and offspring) persist into adulthood. In this study, we used longitudinal data on wild chimpanzees (Pan troglodytes schweinfurthii) at Gombe National Park, Tanzania, to investigate the effects of party-level association during infancy and juvenility (hereafter immaturity) on affiliative relationships among adult males (the philopatric sex). In total, we examined behavioral data from focal follows between 1970 and 2015 (N = 23 adult males). We used maternal associations as proxies for the associations of immature males, and we measured adult male social relationships using party-level associations and grooming activity. We found that immature males that associated with individual adult males at higher rates had stronger relationships with those same adult males later in life. By contrast, rates of association between pairs of immature males did not predict the strength of their dyadic relationships as adults. Overall, these findings emphasize the importance of early socialization in male chimpanzee social development. These results also reinforce studies in mountain gorillas (Gorilla beringei) and bottlenose dolphins (Tursiops sp.), suggesting that the persistence of social relationships that do not involve the mother may be more likely to evolve in long-lived species where young adult males face challenges entering an adult hierarchy composed of stronger and/or more socially experienced competitors. [ABSTRACT FROM AUTHOR]

The mother-offspring relationship is paramount in most mammals and infant survival often depends on maternal investment. In species with prolonged periods of development or co-residence, mothers may continue to influence their offspring's outcomes beyond nutritional independence with benefits biased towards the philopatric sex. Chimpanzees (Pan troglodytes) are long-lived mammals with a protracted period of immaturity during which offspring continue to travel with their mothers. In contrast to most mammals, chimpanzees are also typically male philopatric. Here, we use over 50 years of demographic data from two communities in Gombe National Park, Tanzania, to examine the survival and longevity of both male and female chimpanzees that experienced maternal loss during three different age categories. Males who were orphaned between the ages of 0–4.99 years, 5–9.99 years, and 10–14.99 years all faced significantly lower survival than non-orphans and died earlier than expected. Females faced similarly reduced survival probabilities when orphaned between 0–4.99 and 5–9.99 years of age; however, females who experienced maternal loss between 10 and 14.99 years of age were no more likely to die than non-orphans. Females orphaned in this later age class also lived significantly longer beyond maternal loss than their male counterparts. As observed in other mammals, philopatric male chimpanzees may continue to benefit from their mother's ecological knowledge, whereas maternal influence on female offspring likely fades as they prepare to emigrate. These results emphasize how maternal influence on offspring outcomes can extend well beyond weaning, particularly for the philopatric sex. Significance statement: Mammalian mothers are crucial to their infant's survival and in species where offspring continue to live with their mothers after weaning, maternal influence may extend beyond dependence on mother's milk. While in most group-living mammals females remain in their natal group, chimpanzees typically display the opposite pattern with males residing alongside their mothers into adulthood. Using over 50 years of data on wild chimpanzees, we investigated the consequences of maternal loss both before and after weaning. We found that both males and females orphaned up to 10 years of age were less likely to survive than non-orphans; however, only males orphaned between 10 and 15 years also faced lower survival. These results emphasize how chimpanzee mothers continue to matter for offspring beyond infancy and provide novel data on reduced maternal influence on survival for daughters compared to sons. [ABSTRACT FROM AUTHOR]

Chimpanzees exhibit cultural variation, yet examples of successful cultural transmission between wild communities are lacking. Here we provide the first account of tool-assisted predation ("ant fishing") on Camponotus ants by the Kasekela and Mitumba communities of Gombe National Park. We then consider three hypotheses for the appearance and spread of this behavior in Kasekela: (1) changes in prey availability or other environmental factors, (2) innovation, sod (3) introduction. Ant fishing was recognized as habitual in the Mitumba community by 1992, soon after their habituation began. Apart from one session in 1978, Camponotus predation (typically with tools) was documented in the Kasekela community beginning only in 1994, despite decades of prior observation. By February 2010, ant fishing was customary in Kasekela and with one exception was practiced exclusively by chimpanzees born after 1981 and immigrant females. We hypothesize that changes insect prey availability over time and/or the characteristics of one popular ant-fishing site may have influenced the establishment of ant fishing. Though innovation cannot be completely ruled out, the circumstantial evidence suggests that a Mitumba immigrant introduced ant fishing to Kasekela. We submit that this report represents the first documented case of successful transmission of a novel cultural behavior between wild chimpanzee communities. [ABSTRACT FROM AUTHOR]

Mammalian females generally carry the bulk of reproductive costs. They gestate for relatively long periods of time and provide the majority of parental care for dependent offspring. For this reason, many studies have examined how females deal with the energetic costs of reproduction. Here, we examine the influence of reproductive state on activity budgets, diet quality, and sociality in free-living female chimpanzees (Pan troglodytes) at Gombe National Park, Tanzania. After controlling for dominance rank, we found that pregnant and lactating females consumed higher quality foods than nonpregnant, nonlactating females. However, pregnant females also traveled less. This result did not reflect differences in sociality, as the pregnant female group sizes included in our analyses were comparable to those in other reproductive categories. [ABSTRACT FROM PUBLISHER]

Studies from many different taxa have demonstrated that dominance rank greatly influences individual space use. While the importance of dominance among female chimpanzees, Pan troglodytes, was debated in the past, mounting evidence now shows that rank is very important. In particular, rank has been shown to influence body mass, foraging strategies, association patterns, and ultimately, reproductive success. In this study, we investigated how rank influenced female space use among chimpanzees, P.t. schweinfurthii, at Gombe National Park, Tanzania. Analysing 10 years of data, we found that new immigrants used areas away from dominant females, and that subordinates had lower site fidelity. We also found that high-ranking females had smaller core areas and that this size difference was pronounced during periods of food scarcity when food competition is highest. These patterns suggest that dominant females outcompete subordinates, forcing them to settle elsewhere, range more widely, and shift their space use across time. [Copyright &y& Elsevier]

Researchers have reported a total of 31 infanticides in 4 different chimpanzee (Pan troglodytes) populations. Though infanticide is infrequent, low reproductive rates of females likely make it a strong selective pressure in the species. We report a new incident of intragroup infanticide in Gombe National Park, Tanzania, in which a community male attacked a 3.5-yr-old male. We then consider the infanticide in terms of adaptive and nonadaptive explanations for infanticide including the social pathology, by-product of male aggression, nutritive benefits, resource competition, and sexual selection hypotheses. The incident reported here is not well explained by any of them. While the infanticide is puzzling in terms of ultimate explanations for infanticide, it provides a good context in which to consider proximate mechanisms for offspring recognition. The incident provides some evidence that males may use their mating history with the mother to assess paternity likelihood. [ABSTRACT FROM AUTHOR]

Among mammals, female reproduction is generally thought to be food limited, and dominance should theoretically afford high-ranking females with access to better food resources. Although the importance of dominance rank among female chimpanzees (Pan troglodytes) has been debated in the past, mounting evidence suggests that rank is very important among females (P. t. schweinfurthii) at Gombe National Park, Tanzania. In this study, we investigated the influence of season and dominance rank on female foraging strategies. We found that high-ranking females spent less time foraging and tended to have a narrower diet breadth and higher diet quality than subordinate females. In this way, subordinate female foraging strategies were consistent with how females in general adapted to periods of food scarcity. The results of this study therefore suggest that low-ranking females may face persistent “food scarcity” as a result of interference food competition. We also provide evidence that subordinates may forage less efficiently because they occupy lower quality habitats or avoid associating with dominant females in shared areas. [ABSTRACT FROM AUTHOR]

Although rare among group-living primates, infanticide by females has been reported in several chimpanzee, Pan troglodytes , populations. We examined 13 infanticidal attacks over 47 years at Gombe National Park, Tanzania to evaluate three adaptive hypotheses. (1) Exploitation of the infant as a food resource – by eating a vulnerable neonate, attackers gain calories that may be important during periods of food scarcity or energetic stress. (2) Resource competition – Gombe females concentrate their foraging in overlapping core areas and dominance rank influences foraging success. By killing the infant of a female with high core area overlap, the perpetrator removes a current and future competitor, improving her access to food. (3) Low cost – female chimpanzees mature and reproduce slowly, and longevity increases reproductive success. Physical aggression causes risk of severe injury or death, so females will only mount attacks when risks to the perpetrator are low. In support of hypothesis 1, females usually consumed the carcass. However, attacks were not more likely in times of resource or energy scarcity. In support of hypothesis 2, females attacked others with whom they shared core areas, but attacks did not cause shifts in ranging patterns. In support of hypothesis 3, one or more attackers always outranked the victim, the attacks often involved coalitions and victims usually lacked kin support. Attacks were more likely to be successful when attackers were not hindered by clinging infants and victims could not retreat. Our results provide further evidence for female competition and the adaptive value of female-led infanticide in this species. • We documented 13 female-led infanticidal attacks in one chimpanzee community. • Infant victims were usually eaten, providing immediate nutrition to the attacker. • Attackers targeted likely competitors whose foraging areas overlapped their own. • Attackers usually targeted neonates of lower-ranking mothers who lacked kin support. [ABSTRACT FROM AUTHOR]

A large body of literature demonstrates the adaptive benefits of social relationships between kin, including fitness and survival. Given that most social mammals are characterized by male-biased dispersal, the majority of research on kin selection and associated advantages focuses on social relationships between female kin. Meanwhile, research on social relationships between adult male and female kin has primarily focused on inbreeding avoidance or the benefit to adult sons, with less attention on potential advantages these social relationships may provide females. The general pattern of male dominance over females in most mammal species suggests that females may benefit from protective associations with adult male kin. Using 43 years of behavioural data on the wild chimpanzees, Pan troglodytes schweinfurthii , of Gombe National Park, Tanzania, we examined association patterns between females and their adult maternal male kin. We specifically focused on how these associations may represent a trade-off between inbreeding avoidance and protection for females, particularly against infanticide. In accordance with inbreeding avoidance, we predicted that females' association with adult kin would decrease when they were maximally tumescent, signalling sexual receptivity. To determine whether female–male kin associations provide protection to females, we examined female associations with adult male kin during their first year postpartum when infants are most vulnerable to infanticide. We predicted that during this first year postpartum, females would have a higher association with male kin than with unrelated males. We found that females associated more with adult sons and brothers than with unrelated males when they did not have a sexual swelling. Female association increased with all males across tumescence but females associated less with their brothers than they did with their sons and unrelated males when they were maximally tumescent, providing equivocal support for the inbreeding avoidance hypothesis. Furthermore, females associated more with both sons and brothers than with unrelated males in the first 6 months of the postpartum period. Higher association with brothers, relative to unrelated males, persisted throughout the first year postpartum. Together, these results speak to the cost–benefit trade-off in female and adult male kin associations, highlighting the potential protective advantage for females, especially during the postpartum period. • We examined association patterns between females and their adult maternal male kin. • Association with sons, brothers and unrelated males increased with swelling size. • Association with sons was higher than with other males during early postpartum. • Association with brothers was higher than with unrelated males 1 year postpartum. • Our results suggest a potential male kin protective advantage postpartum. [ABSTRACT FROM AUTHOR]

Objectives: Many nonhuman primate diets are dominated by plant foods, yet plant tissues are often poor sources of sodium-a necessary mineral for metabolism and health. Among primates, chimpanzees (Pan troglodytes), which are ripe fruit specialists, consume diverse animal, and plant resources. Insects have been proposed as a source of dietary sodium for chimpanzees, yet published data on sodium values for specific foods are limited. We assayed plants and insects commonly eaten by chimpanzees to assess their relative value as sodium sources., Materials and Methods: We used atomic absorption spectroscopy to determine sodium content of key plant foods and insects consumed by chimpanzees of Gombe National Park, Tanzania. Dietary contributions of plant and insect foods were calculated using feeding observational data., Results: On a dry matter basis, mean sodium value of plant foods (n = 83 samples; mean = 86 ppm, SD = 92 ppm) was significantly lower than insects (n = 12; mean = 1549 ppm, SD = 807 ppm) (Wilcoxon rank sum test: W = 975, p < 0.001). All plant values were below the suggested sodium requirement (2000 ppm) for captive primates. While values of assayed insects were variable, sodium content of two commonly consumed insect prey for Gombe chimpanzees (Macrotermes soldiers and Dorylus ants) were four to five times greater than the highest plant values and likely meet requirements., Discussion: We conclude that plant foods available to Gombe chimpanzees are generally poor sources of sodium while insects are important, perhaps critical, sources of sodium for this population., (© 2024 Wiley Periodicals LLC.)

Objectives: A key feature of human life history evolution is that modern humans wean their infants 2-4 years earlier on average than African apes. However, our understanding of weaning variation in apes remains limited. Here we provide the first such report in chimpanzees by examining weaned age variation using long-term data from Gombe National Park, Tanzania., Material and Methods: We analyzed 41 years of observational behavioral data from 65 offspring of 29 mothers to examine the relationships between weaned age (defined as cessation of suckling) in wild chimpanzees and maternal age, dominance rank and parity, and offspring sex. We used Cox proportional hazards regression with mixed effects to model time to weaning and to examine potential sources of variation in offspring weaned age., Results: We found that male offspring were less likely than female offspring to wean by a given age and that weaned age of males varied more than weaned age of females. In addition, maternal dominance rank interacted with offspring age, such that low-ranking mothers were less likely to wean offspring early, but this effect decreased with offspring age., Discussion: We found that male offspring and offspring of low-ranking females were less likely to wean early, but did not find evidence for variable weaning according to maternal age or parity. As more data accumulate, we will be better able to disentangle the effects of maternal dominance rank, age and parity. Such studies will not only provide a richer understanding of living ape life history characteristics, but will also provide an important framework for understanding the evolution of early weaning in humans., (© 2019 Wiley Periodicals, Inc.)

Objectives: Primates exhibit variation in rates of growth and development. Variation in female growth and development across ape species appears to be explained by the Ecological Risk Aversion Hypothesis (ERAH). Indeed, existing data on variation in somatic growth and reproductive maturation between humans' closest living ape relatives, bonobos and chimpanzees, appear to be consistent with this hypothesis. However, existing data on behavioral maturation between the two species appear to contradict this hypothesis. We present novel behavioral data on infant and juvenile females from wild populations of both species in order to further evaluate predictions of the ERAH as it relates to the speed of behavioral maturation., Materials and Methods: We analyzed 3 years of behavioral data on 17 female bonobos (<8 years of age) from LuiKotale, Democratic Republic of the Congo and 40 years of behavioral data on 30 age-matched female chimpanzees from Gombe, Tanzania. We compared the timing of (a) the attainment of independence from mothers and (b) the development of social skills using the following proxies: proximity between females and their mothers and the time that females spent engaged in eating, suckling, social play, social grooming, and riding on their mothers., Results: We did not find species differences in the proportion of time that females spent in contact with their mothers or engaged in eating, suckling, social play, or social grooming. Female bonobos spent more time riding on their mothers than did female chimpanzees. Female bonobos spent more time at distances greater than 5 m from their mothers during the ages of 3-8 years, but females did not differ during the ages of 0-3 years., Discussion: Behavioral maturation is largely similar between females of the two species based on the ages and proxies considered herein. We propose alternative explanations for the differences that we found in proximity and riding that do not invoke differences in underlying rates of maturation., (© 2019 Wiley Periodicals, Inc.)

Early life experiences are known to influence hypothalamic-pituitary-adrenal (HPA) axis development, which can impact health outcomes through the individual's ability to mount appropriate physiological reactions to stressors. In primates, these early experiences are most often mediated through the mother and can include the physiological environment experienced during gestation. Here, we investigate stress physiology of dependent offspring in wild chimpanzees for the first time and examine whether differences in maternal stress physiology are related to differences in offspring stress physiology. Specifically, we explore the relationship between maternal rank and maternal fecal glucocorticoid metabolite (FGM) concentration during pregnancy and early lactation (first 6 months post-partum) and examine whether differences based on maternal rank are associated with dependent offspring FGM concentrations. We found that low-ranking females exhibited significantly higher FGM concentrations during pregnancy than during the first 6 months of lactation. Furthermore, during pregnancy, low-ranking females experienced significantly higher FGM concentrations than high-ranking females. As for dependent offspring, we found that male offspring of low-ranking mothers experienced stronger decreases in FGM concentrations as they aged compared to males with high-ranking mothers or their dependent female counterparts. Together, these results suggest that maternal rank and FGM concentrations experienced during gestation are related to offspring stress physiology and that this relationship is particularly pronounced in males compared to females. Importantly, this study provides the first evidence for maternal effects on the development of offspring HPA function in wild chimpanzees, which likely relates to subsequent health and fitness outcomes. Am. J. Primatol. 80:e22525, 2018. © 2016 Wiley Periodicals, Inc., (© 2016 Wiley Periodicals, Inc.)

Disease and other health hazards pose serious threats to the persistence of wild ape populations. The total chimpanzee population at Gombe National Park, Tanzania, has declined from an estimated 120 to 150 individuals in the 1960's to around 100 individuals by the end of 2013, with death associated with observable signs of disease as the leading cause of mortality. In 2004, we began a non-invasive health-monitoring program in the two habituated communities in the park (Kasekela and Mitumba) with the aim of understanding the prevalence of health issues in the population, and identifying the presence and impacts of various pathogens. Here we present prospectively collected data on clinical signs (observable changes in health) in the chimpanzees of the Kasekela (n = 81) and Mitumba (n = 32) communities over an 8-year period (2005-2012). First, we take a population approach and analyze prevalence of clinical signs in five different categories: gastrointestinal system (diarrhea), body condition (estimated weight loss), respiratory system (coughing, sneezing etc.), wounds/lameness, and dermatologic issues by year, month, and community membership. Mean monthly prevalence of each clinical sign per community varied, but typically affected <10% of observed individuals. Secondly, we analyze the presence of clinical signs in these categories as they relate to individual demographic and social factors (age, sex, and dominance rank) and simian immunodeficiency virus (SIVcpz) infection status. Adults have higher odds of being observed with diarrhea, loss of body condition, and wounds or lameness when compared to immatures, while males have a higher probability of being observed with wounds or lameness than females. In contrast, signs of respiratory illness appear not to be related to chimpanzee-specific factors and skin abnormalities are very rare. For a subset of known-rank individuals, dominance rank predicts the probability of wounding/lameness in adult males, but does not predict any adverse clinical signs in adult females. Instead, adult females with SIVcpz infection are more likely to be observed with diarrhea, a finding that warrants further investigation. Comparable data are needed from other sites to determine whether the prevalence of clinical signs we observe are relatively high or low, as well as to more fully understand the factors influencing health of wild apes at both the population and individual level. Am. J. Primatol. 80:e22562, 2018. © 2016 Wiley Periodicals, Inc., (© 2016 Wiley Periodicals, Inc.)

Immunoglobulin A (IgA) is the primary antibody responsible for mucosal defense in mammals and has been used as a marker for chronic stress and immune status. Therefore, this antibody may provide a more reliable indicator of an individual's immunocompetence than is currently available through other methods. Immunoglobulin A has never before been quantified in a wild population of non-human primates using non-invasive sample collection techniques. In this study, we present methodology for non-invasive IgA extraction in the field and provide quantification of mean fecal IgA concentrations in wild chimpanzees (Pan troglodytes schweinfurthii). During the study period (November 2009-October 2010), we collected fecal samples (N = 1463) from 59 individuals at Gombe National Park, Tanzania. We modified a field extraction technique for steroidal hormones to extract IgA from the fecal samples and then quantified mean IgA concentrations (ng/g) using a commercial human IgA enzyme immunoassay. Mean IgA concentration varied among individuals but not by sex or reproductive status. Mature animals tended toward higher mean IgA concentration than immature. Mean IgA concentration differed by quartile season, following a similar pattern previously observed for respiratory illness rates in this population, with the late dry season having significantly higher averages than the late wet. A circadian rhythm was also evident with mean IgA concentrations higher in samples collected in the latter half of the day. These demographic and temporal patterns of IgA concentration provide baseline values necessary to interpret future results, which may be combined with other health values to better understand the role of health and long-term stress in wild great ape populations. Am. J. Primatol. 80:e22558, 2018. © 2016 Wiley Periodicals, Inc., (© 2016 Wiley Periodicals, Inc.)

The role of biological and social influences on sex differences in human child development is a persistent topic of discussion and debate. Given their many similarities to humans, chimpanzees are an important study species for understanding the biological and evolutionary roots of sex differences in human development. In this study, we present the most detailed analyses of wild chimpanzee infant development to date, encompassing data from 40 infants from the long-term study of chimpanzees at Gombe National Park, Tanzania. Our goal was to characterize age-related changes, from birth to five years of age, in the percent of observation time spent performing behaviors that represent important benchmarks in nutritional, motor, and social development, and to determine whether and in which behaviors sex differences occur. Sex differences were found for indicators of social behavior, motor development and spatial independence with males being more physically precocious and peaking in play earlier than females. These results demonstrate early sex differentiation that may reflect adult reproductive strategies. Our findings also resemble those found in humans, which suggests that biologically-based sex differences may have been present in the common ancestor and operated independently from the influences of modern sex-biased parental behavior and gender socialization.

Monitoring adrenocortical activity in wild primate populations is critical, given the well-documented relationship between stress, health, and reproduction. Although many primate studies have quantified fecal glucocorticoid metabolite (FGM) concentrations, it is imperative that researchers validate their method for each species. Here, we describe and validate a technique for field extraction and storage of FGMs in wild chimpanzees (Pan troglodytes). Our method circumvents many of the logistical challenges associated with field studies while yielding similar results to a commonly used laboratory method. We further validate that our method accurately reflects stress physiology using an adrenocorticotropic hormone challenge in a captive chimpanzee and an FGM peak at parturition in a wild subject. Finally, we quantify circadian patterns for FGMs for the first time in this species. Understanding these patterns may allow researchers to directly link specific events with the stress response., (© 2012 Wiley Periodicals, Inc.)

Infectious disease and other health hazards have been hypothesized to pose serious threats to the persistence of wild ape populations. Respiratory disease outbreaks have been shown to be of particular concern for several wild chimpanzee study sites, leading managers, and researchers to hypothesize that diseases originating from and/or spread by humans pose a substantial risk to the long-term survival of chimpanzee populations. The total chimpanzee population in Gombe National Park, Tanzania, has declined from 120-150 in the 1960s to about 100 by the end of 2007, with death associated with observable signs of disease as the leading cause of mortality. We used a historical data set collected from 1979 to 1987 to investigate the baseline rates of respiratory illness in chimpanzees at Gombe National Park, Tanzania, and to analyze the impact of human-related factors (e.g., banana feeding, visits to staff quarters) and non-human-related factors (e.g., sociality, season) on chimpanzee respiratory illness rates. We found that season and banana feeding were the most significant predictors of respiratory health clinical signs during this time period. We discuss these results in the context of management options for the reduction of disease risk and the importance of long-term observational data for conservation.