RESUMEN
Maintaining water balance is essential for organismal health, and lactating females must balance individual needs with milk production and offspring hydration. Primate milk is dilute and presumed to be the primary source for infant hydration for a considerable time period. Few studies have investigated the hydration burden that lactation may place on female primates. In this study, we investigated sources of variation in female and offspring drinking frequency among wild chimpanzees (Pan troglodytes). We hypothesized females would experience seasonal and lactation hydration burdens and adjust their drinking behavior to accommodate these, but this hydration burden would vary between females of different dominance ranks. We also predicted that parity would relate to maternal drinking frequency since primiparous females are still investing in their own growth. Finally, we predicted that offspring would drink more in the dry season and as they aged and lost milk as a water source, but that offspring of high-ranking females would be buffered from these effects. Using 41 years of long-term data on the behavior of mothers and offspring of Gombe National Park, we found that mothers drank more in the dry season, but there was no significant difference between mothers of different ranks during this period. Low-ranking females drank significantly more than mid- and high-ranking females during late lactation. Offspring also drank more in the dry season and as they aged, but there was no evidence of buffering for those with high-ranking mothers. While chimpanzees in our study population drank infrequently, they do demonstrate noticeable shifts in drinking behavior that suggests seasonal and reproductive hydration burdens.
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Lactancia , Pan troglodytes , Animales , Femenino , Humanos , Madres , Embarazo , Reproducción , AguaRESUMEN
Infectious disease outbreaks pose a significant threat to the conservation of chimpanzees (Pan troglodytes) and all threatened nonhuman primates. Characterizing and mitigating these threats to support the sustainability and welfare of wild populations is of the highest priority. In an attempt to understand and mitigate the risk of disease for the chimpanzees of Gombe National Park, Tanzania, we initiated a long-term health-monitoring program in 2004. While the initial focus was to expand the ongoing behavioral research on chimpanzees to include standardized data on clinical signs of health, it soon became evident that the scope of the project would ideally include diagnostic surveillance of pathogens for all primates (including people) and domestic animals, both within and surrounding the National Park. Integration of these data, along with in-depth post-mortem examinations, have allowed us to establish baseline health indicators to inform outbreak response. Here, we describe the development and expansion of the Gombe Ecosystem Health project, review major findings from the research and summarize the challenges and lessons learned over the past 16 years. We also highlight future directions and present the opportunities and challenges that remain when implementing studies of ecosystem health in a complex, multispecies environment.
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Ecosistema , Pan troglodytes , Animales , Humanos , Estudios Longitudinales , Parques Recreativos , Primates , Tanzanía/epidemiologíaRESUMEN
OBJECTIVES: The prolonged juvenile period exhibited by primates is an evolutionary conundrum. Here we examine wild chimpanzee feeding development in the context of two hypotheses regarding prolonged development in primates: the needing-to-learn hypothesis and the expensive brain hypothesis. MATERIAL AND METHODS: We studied wild chimpanzee (Pan troglodytes schweinfurthii) offspring at Gombe National Park, Tanzania. We analyzed 41 years of observational behavioral data collected between 1975 and 2016 from 81 offspring. We characterized feeding development in the first 10 years of life via four different measures: (1) proportion of observation time spent feeding; (2) diet composition; (3) diet breadth; and (4) diet maturity as measured by similarity to maternal diet. We used mixed effects models to examine changes with age and by sex, while controlling for season. RESULTS: Feeding time, diet breadth, and diet maturity exhibited the most substantial increases with age in the first 6 years, with no significant change thereafter. Males and females showed different patterns of change in diet breadth by age, but did not differ by age 10. Diet composition did not change significantly with age and did not differ by sex. DISCUSSION: We found that chimpanzee offspring attained adult-like feeding behaviors between 4 and 6 years of age, concomitant with the completion of weaning. Thus, our data do not support the needing-to-learn feeding skills hypothesis of a prolonged juvenile period, but additional data are needed to evaluate how and when adolescent chimpanzees are able to make foraging decisions independent of their mothers. Existing data on growth provides support for the expensive brain hypothesis, however, these hypotheses are not necessarily mutually exclusive. As more studies across taxa accumulate sufficient datasets on a range of developmental metrics, we will be able to achieve a more robust understanding of prolonged development in primates.
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Conducta Alimentaria/fisiología , Pan troglodytes/fisiología , Destete , Animales , Antropología Física , Femenino , Masculino , MadresRESUMEN
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.
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Hominidae , Pan troglodytes , Animales , Humanos , Pan paniscus , Tanzanía , Erupción DentalRESUMEN
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.
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Pan troglodytes/fisiología , Predominio Social , Destete , Factores de Edad , Animales , Femenino , Masculino , Modelos Biológicos , Parques Recreativos , TanzaníaRESUMEN
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.
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Conducta Animal , Pan paniscus/psicología , Pan troglodytes/psicología , Conducta Social , Animales , República Democrática del Congo , Femenino , Pan paniscus/crecimiento & desarrollo , Pan troglodytes/crecimiento & desarrollo , TanzaníaRESUMEN
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.
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Heces/química , Inmunoglobulina A/análisis , Pan troglodytes , Hormona Adrenocorticotrópica/administración & dosificación , Envejecimiento , Animales , Ritmo Circadiano/fisiología , Ensayo de Inmunoadsorción Enzimática/métodos , Ensayo de Inmunoadsorción Enzimática/veterinaria , Femenino , Inmunoglobulina A/efectos de los fármacos , Masculino , Reproducción/fisiología , Estaciones del Año , TanzaníaRESUMEN
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.
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Sistema Hipotálamo-Hipofisario/fisiología , Pan troglodytes/fisiología , Sistema Hipófiso-Suprarrenal/fisiología , Estrés Fisiológico/fisiología , Envejecimiento , Animales , Conducta Animal/fisiología , Heces/química , Femenino , Glucocorticoides/análisis , Glucocorticoides/metabolismo , Lactancia/fisiología , Masculino , Pan troglodytes/psicología , Embarazo/fisiología , Efectos Tardíos de la Exposición Prenatal , Predominio Social , TanzaníaRESUMEN
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.
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Estado de Salud , Pan troglodytes , Predominio Social , Factores de Edad , Animales , Diarrea/veterinaria , Estudios Longitudinales , Pan troglodytes/lesiones , Prevalencia , Enfermedades Respiratorias/veterinaria , Factores Sexuales , Síndrome de Inmunodeficiencia Adquirida del Simio/epidemiología , Enfermedades de la Piel/veterinaria , Tanzanía , Pérdida de PesoRESUMEN
Compared to great apes, humans maintain a relatively rapid reproductive pace despite long periods of dependency. This seemingly contradictory set of traits is made possible by weaning offspring before nutritional independence and alloparents who help provide care. In traditional societies, this help may be provided to mothers in part by their juvenile offspring who carry, supervise, or provision younger siblings. In contrast to humans, chimpanzees (Pan troglodytes) are nutritionally independent after weaning, yet juveniles continue to travel with their mother and younger sibling for an additional 4-5 years. This continued association could be costly to the mother if she continues to invest in weaned offspring. Alternately, while juvenile chimpanzees do not typically provision younger siblings, their presence and social interaction with infants may allow mothers to focus on other tasks. In this study, we investigate the costs and benefits to mothers of continued association with juveniles in wild chimpanzees. Using 26 years of long-term behavioral data we examined how maternal activity budgets varied based on the presence of a dependent juvenile offspring. We found that continued social interaction between mothers and juveniles does not influence the mother's time allocated to interacting with the younger infant, her feeding, resting, or travel time, or time socializing with other community members. Instead, mothers may benefit from the additional social interaction and/or relationship with their older offspring. Using 45 years of demographic data we found that those offspring who had an older sibling tended to be more likely to survive each year from birth to 8 years than those without an older sibling. Additionally, interbirth intervals were more likely to end when the female had an older offspring present. A mutually beneficial mother-juvenile dynamic in great apes provides insight into continued association between mothers and offspring after nutritional independence and the emergence of juvenile helping during hominin evolution.
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Composición Familiar , Conducta Materna , Madres , Pan troglodytes/fisiología , Pan troglodytes/psicología , Reproducción , Destete , Factores de Edad , Animales , FemeninoRESUMEN
Among modern foraging societies, men hunt more than women, who mostly target relatively low-quality, reliable resources (i.e., plants). This difference has long been assumed to reflect human female reproductive constraints, particularly caring for and provisioning mates and offspring. Long-term studies of chimpanzees (Pan troglodytes) enable tests of hypotheses about the possible origins of human sex differences in hunting, prior to pair-bonding and regular provisioning. We studied two eastern chimpanzee communities (Kasekela, Mitumba) in Gombe, Tanzania and one (Kanyawara) in Kibale, Uganda. Relative to males, females had low hunting rates in all three communities, even where they encountered red colobus monkeys (the primary prey of chimpanzees) as often as males did. There was no evidence that clinging offspring hampered female hunting. Instead, consistent with the hypothesis that females should be more risk-averse than males, females at all three sites specialized in low-cost prey (terrestrial/sedentary prey at Gombe; black and white colobus monkeys at Kanyawara). Female dominance rank was positively correlated with red colobus hunting probability only at Kasekela, suggesting that those in good physical condition were less sensitive to the costs of possible failure. Finally, the potential for carcass appropriation by males deterred females at Kasekela (but not Kanyawara or Mitumba) from hunting in parties containing many adult males. Although chimpanzees are not direct analogs of the last common ancestor (LCA) of Pan and Homo, these results suggest that before the emergence of social obligations regarding sharing and provisioning, constraints on hunting by LCA females did not necessarily stem from maternal care. Instead, they suggest that a risk-averse foraging strategy and the potential for losing prey to males limited female predation on vertebrates. Sex differences in hunting behavior would likely have preceded the evolution of the sexual division of labor among modern humans.
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Carne , Pan troglodytes , Conducta Predatoria , Factores Sexuales , Animales , Colobus , Femenino , Hominidae , Masculino , Caracteres Sexuales , Tanzanía , UgandaRESUMEN
Social play is common among many group-living animals, but the benefits are not well understood. Proposed benefits include increased muscle coordination as the result of increased locomotor versatility and development, and strengthened social bonds through interactions with like-aged individuals. In this study, we used 33 years of long-term behavioral data on infant chimpanzees in Gombe National Park, Tanzania, to examine these potential benefits of social play, specifically how the percentage of time engaged in social play relates to motor and social developmental milestones. We predicted that infants who engaged in more social play would achieve motor and social milestones at younger ages. We found that individuals that spent more time engaging in social play achieved the motor milestones of riding dorsally and traveling independently at earlier ages. Additionally, we found that the amount of play was correlated with earlier ages for reaching the social milestones of spatial independence from mother, first grooming of non-maternal kin, and first observed mating attempt. This is the first study in great apes to demonstrate a relationship between play behavior and developmental milestones, supporting the hypotheses that play provides motor, and social benefits.
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Actividad Motora , Pan troglodytes/fisiología , Conducta Social , Animales , Femenino , Masculino , Pan troglodytes/crecimiento & desarrollo , Juego e Implementos de Juego , TanzaníaRESUMEN
In many mammals, early social experience is critical to developing species-appropriate adult behaviors. Although mother-infant interactions play an undeniably significant role in social development, other individuals in the social milieu may also influence infant outcomes. Additionally, the social skills necessary for adult success may differ between the sexes. In chimpanzees (Pan troglodytes), adult males are more gregarious than females and rely on a suite of competitive and cooperative relationships to obtain access to females. In fission-fusion species, including humans and chimpanzees, subgroup composition is labile and individuals can vary the number of individuals with whom they associate. Thus, mothers in these species have a variety of social options. In this study, we investigated whether wild chimpanzee maternal subgrouping patterns differed based on infant sex. Our results show that mothers of sons were more gregarious than mothers of daughters; differences were especially pronounced during the first 6 mo of life, when infant behavior is unlikely to influence maternal subgrouping. Furthermore, mothers with sons spent significantly more time in parties containing males during the first 6 mo. These early differences foreshadow the well-documented sex differences in adult social behavior, and maternal gregariousness may provide sons with important observational learning experiences and social exposure early in life. The presence of these patterns in chimpanzees raises questions concerning the evolutionary history of differential social exposure and its role in shaping sex-typical behavior in humans.
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Animales Salvajes/fisiología , Conducta Animal , Pan troglodytes/fisiología , Conducta Social , Animales , Femenino , MasculinoRESUMEN
An increase in faunivory is a consistent component of human evolutionary models. Animal matter is energy- and nutrient-dense and can provide macronutrients, minerals, and vitamins that are limited or absent in plant foods. For female humans and other omnivorous primates, faunivory may be of particular importance during the costly periods of pregnancy and early lactation. Yet, because animal prey is often monopolizable, access to fauna among group-living primates may be mediated by social factors such as rank. Wild chimpanzees (Pan troglodytes) across Africa habitually consume insects and/or vertebrates. However, no published studies have examined patterns of female chimpanzee faunivory during pregnancy and early lactation relative to non-reproductive periods, or by females of different rank. In this study, we assessed the influence of reproductive state and dominance rank on the consumption of fauna (meat and insects) by female chimpanzees of Gombe National Park, Tanzania. Using observational data collected over 38 years, we tested (a) whether faunivory varied by reproductive state, and (b) if high-ranking females spent more time consuming fauna than lower-ranking females. In single-factor models, pregnant females consumed more meat than lactating and baseline (meaning not pregnant and not in early lactation) females, and high-ranking females consumed more meat than lower-ranking females. A two-factor analysis of a subset of well-sampled females identified an interaction between rank and reproductive state: lower-ranking females consumed more meat during pregnancy than lower-ranking lactating and baseline females did. High-ranking females did not significantly differ in meat consumption between reproductive states. We found no relationships between rank or reproductive state with insectivory. We conclude that, unlike insectivory, meat consumption by female chimpanzees is mediated by both reproductive state and social rank. We outline possible mechanisms for these patterns, relate our findings to meat-eating patterns in women from well-studied hunter-gatherer societies, and discuss potential avenues for future research.
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Conducta Alimentaria/fisiología , Carne , Pan troglodytes/fisiología , Reproducción/fisiología , Predominio Social , Animales , FemeninoRESUMEN
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.
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Pan troglodytes , Animales , Pan troglodytes/metabolismo , Tanzanía , Dieta/veterinaria , Insectos , Sodio en la Dieta/análisis , Plantas/química , Plantas/metabolismo , Sodio/análisis , Conducta Alimentaria/fisiología , Femenino , Alimentación Animal/análisis , MasculinoRESUMEN
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.
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Heces/química , Glucocorticoides/metabolismo , Pan troglodytes/fisiología , Hormona Adrenocorticotrópica , Animales , Ritmo Circadiano , Femenino , Técnicas para Inmunoenzimas , Inyecciones Intramusculares , Masculino , Parto , Estaciones del Año , Estrés Fisiológico , TanzaníaRESUMEN
Space use often correlates with reproductive success [1, 2]. Individual site fidelity is ubiquitous across a variety of taxa, including birds, mammals, insects, and reptiles [3-9]. Individuals can benefit from using the same area because doing so affords access to known resources, including food and/or breeding sites. The majority of studies on site fidelity have focused upon strictly territorial species in which individuals range in well-defined, exclusive areas (e.g., [4, 9]). By comparison, the transient groups that define fission-fusion species allow for considerable flexibility in individual space use. Although there is evidence that individual space use can influence reproductive success [2], relatively little is known about individual ranging patterns in fission-fusion species. Here, we investigate three potential correlates of male site fidelity (age, habitat quality, and maternal space use) in wild chimpanzees (Pan troglodytes). We found that when alone, each male preferentially concentrated his space use near the area where his mother ranged when he was dependent. We suggest that solitary ranging allows males to avoid direct competition with conspecifics and that foraging in familiar areas maximizes foraging efficiency. These results highlight the importance of male foraging strategies in a species in which male ranging is typically explained in terms of mating access to females.
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Fenómenos de Retorno al Lugar Habitual , Pan troglodytes/fisiología , Factores de Edad , Animales , Ambiente , Femenino , Masculino , Factores SexualesRESUMEN
Fission-fusion dynamics have evolved in a broad range of animal taxa and are thought to allow individuals to mitigate feeding competition. While this is the principal benefit of fission-fusion, few studies have evaluated its costs. We compared gregariousness, foraging budgets, and social budgets between lactating bonobos and chimpanzees from wild populations to evaluate potential costs. Both species exhibit fission-fusion dynamics, but chimpanzees, particularly in East African populations, appear to experience higher feeding competition than bonobos. We expected lactating chimpanzees to be less gregarious than lactating bonobos; reduced gregariousness should allow lactating chimpanzees to mitigate the costs of higher feeding competition without requiring more foraging effort. However, we expected the reduced gregariousness of lactating chimpanzees to limit their time available for affiliative interactions. Using long-term data from LuiKotale bonobos and Gombe chimpanzees, we found that lactating chimpanzees were indeed less gregarious than lactating bonobos, while feeding and travel time did not differ between species. Contrary to our predictions, lactating females did not differ in social interaction time, and lactating chimpanzees spent proportionately more time interacting with individuals other than their immature offspring. Our results indicate that lactating chimpanzees can maintain social budgets comparable to lactating bonobos despite reduced gregariousness and without incurring additional foraging costs. We discuss potential explanations for why lactating bonobos are more gregarious.
RESUMEN
1. Fertility is an important fitness component, but is difficult to measure in slowly reproducing, long-lived animals such as chimpanzees (Pan troglodytes). 2. We measured fertility and the effect of measured covariates on fertility in a 43-year sample of birth intervals of chimpanzees from the Gombe National Park, Tanzania using Cox proportional hazards regression with individual-level random effects. 3. The birth hazard declined with mothers' age at a rate of 0·84 per year following age at first reproduction. This value is somewhat stronger than previous estimates. 4. Loss of the infant that opened the birth interval increased the birth hazard 134-fold. 5. Birth intervals following the first complete birth interval were shorter than this first interval, while sex of the previous infant had no significant effect. 6. Maternal dominance rank was significant at the P < 0·1 level when coded as high/middle/low but was highly significant when we simply considered high rank vs. others. 7. Individual heterogeneity had a substantial impact on birth interval duration. We interpret this individual effect as a measure of phenotypic quality, controlling for the measured covariates such as dominance rank. This interpretation is supported by the correlation of individual heterogeneity scores with similar independent measures of body mass.
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Fertilidad , Pan troglodytes/fisiología , Envejecimiento , Animales , Peso Corporal , Femenino , Masculino , Predominio SocialRESUMEN
The study of non-human primate thanatology has expanded dramatically in recent years as scientists seek to understand the evolutionary roots of human death concepts and practices. However, observations of how conspecifics respond to dead individuals are rare and highly variable. Mothers of several species of primate have been reported to carry and continue to interact with dead infants. Such interactions have been proposed to be related to maternal condition, attachment, environmental conditions or reflect a lack of awareness that the infant has died. Here, we tested these hypotheses using a dataset of cases of infant corpse carrying by chimpanzees in Gombe National Park, Tanzania (n = 33), the largest dataset of such cases in chimpanzees. We found that mothers carried infant corpses at high rates, despite behavioural evidence that they recognize that death has occurred. Median duration of carriage was 1.83 days (interquartile range = 1.03-3.59). Using an information theoretic approach, we found no support for any of the leading hypotheses for duration of continued carriage. We interpret these data in the context of recent discussions regarding what non-human primates understand about death.