Pan Species Research Articles

Does green mean clean? Volatile organic emissions from regular versus green cleaning products.

Cleaning products emit a range of volatile organic compounds (VOCs), including some which are hazardous or can undergo chemical transformations to generate harmful secondary pollutants. In recent years, "green" cleaners have become increasingly popular, with an implicit assumption that these are better for our health and/or the environment. However, there is no strong evidence to suggest that they are better for indoor air quality compared to regular products. In this study, the VOC composition of 10 regular and 13 green cleaners was examined by headspace analysis. Monoterpenes were the most prevalent VOCs, with average total monoterpene concentrations of 8.6 and 25.0 mg L-1 for regular and green cleaners, respectively. Speciated monoterpene emissions were applied to a detailed chemical model to investigate the indoor air chemistry following a typical cleaning event. Green cleaners generally emitted more monoterpenes than regular cleaners, resulting in larger increases in harmful secondary pollutant concentrations following use, such as formaldehyde (up to 7%) and PAN species (up to 6%). However, emissions of the most reactive monoterpenes (α-terpinene, terpinolene and α-phellandrene), were observed more frequently from regular cleaners, resulting in a disproportionately large impact on the concentrations of radical species and secondary pollutants that were formed after cleaning occurred.

Tool use, or not tool use, that is the question: is the necessity hypothesis really inconsequential for the African great apes?

Investigating the drivers of tool use in animals has recently received great attention because of its implication in understanding animals’ cognition and the evolution of tool use in hominins. The necessity hypothesis posits tool use as a necessary response to food scarcity, but its role is an ongoing debate. The largest body of literature comparing animal tool use frequencies is with regard to primates, particularly comparisons between the Pan species. This supports the hypothesis that tool use is rarer in wild bonobos because of differential manipulation abilities of chimpanzees rather than different ecological needs. In this article, I aim to enrich the discussion concerning the necessity hypothesis and the ecological drivers of tool use in apes. The higher feeding flexibility of bonobos may be a key aspect to explaining the lower use of feeding tools than that observed in chimpanzees. The diet flexibility of bonobos is similar to that of the lowest level of tool users among the wild great apes: the gorilla. Gorillas can thus help to shed further light on this debate. When fruit is scarce, Western gorillas and bonobos rely more on widely available proteinaceous herbs than chimpanzees, who remain highly frugivorous. Chimpanzees may thus face a greater necessity to search for an alternative to obtain high-quality food: tool-assisted feeding. An indirect piece of evidence for this higher level of herbivory is that the prevalence of gut ciliates in bonobos is double that of chimpanzees. In each animal species, a different combination of necessity, opportunities, predisposition, and learning processes are likely to be at play in the emergence of flexible tool use in animals.

Ape femoral-humeral rigidities and arboreal locomotion.

This study investigates patterns of bone functional adaptations in extant apes through comparing hindlimb to forelimb bone rigidity ratios in groups with varying levels of arboreality. Using CT scans, bone rigidity (J) was calculated at three regions of interest (ROI) along femoral and humeral diaphyses in Homo, Pongo, Pan, and Gorilla with further comparisons made between species and subspecies divisions within Pan and Gorilla. Consistent with previous work on extant hominoids, species exhibited differences in midshaft femoral to humeral (F/H) rigidity ratios. Results of the present study confirm that these midshaft differences extend to 35% and 65% diaphyseal ROIs. Modern humans, exhibiting larger ratios, and orangutans, exhibiting smaller ratios, bracketed the intermediate African apes in comparisons. Within some African apes, limb rigidity ratios varied significantly between taxonomic groups. Eastern gorillas exhibited the highest mean ratios and chimpanzees the lowest at all three ROIs. In posthoc comparisons, chimpanzees and bonobos did not differ in relative limb rigidity ratios at any of the three ROIs. However, western gorillas were more similar to bonobos than eastern gorillas at 50% and 35% ROIs, but not at the 65% ROI. Species, and to a lesser extent subspecies, can be distinguished by F/H limb rigidity ratios according to broad positional behavior patterns at multiple regions of interest along the diaphyses. Similarity of bonobos and western gorillas is in line with behavioral data of bonobos being the most terrestrial of Pan species, and western gorillas the most arboreal of the Gorilla groups.

Oxytocin and social gaze during a dominance categorization task in tufted capuchin monkeys.

Visual attention to facial features is an important way that group-living primate species gain knowledge about others. However, where this attention is focused on the face is influenced by contextual and social features, and emerging evidence in Pan species suggests that oxytocin, a hormone involved in forming and maintaining affiliative bonds among members of the same group, influences social attention as measured by eye gaze. Specifically, bonobos tend to focus on conspecifics’ eyes when viewing two-dimensional images, whereas chimpanzees focus more on the edges of the face. Moreover, exogenous oxytocin, which was hypothesized to increase eye contact in both species, instead enhanced this existing difference. We follow up on this to (1) determine the degree to which this Pan pattern generalizes across highly social, cooperative non-ape primates and (2) explore the impact of exogenously administered vs. endogenously released oxytocin in impacting this behavior. To do so, we tracked gaze direction on a computerized social categorization task using conspecific faces in tufted capuchin monkeys (Sapajus [Cebus] apella) after (1) exogenously administering intranasal oxytocin using a nebulizer or (2) inducing an endogenous increase in oxytocin using fur-rubbing, previously validated to increase oxytocin in capuchins. Overall, we did not find a general tendency in the capuchins to look toward the eyes or mouth, but we found that oxytocin was related to looking behavior toward these regions, albeit not in a straightforward way. Considering frequency of looking per trial, monkeys were more likely to look at the eye region in the fur-rubbing condition as compared to either the saline or exogenous oxytocin conditions. However, in terms of duration of looking during trials in which they did look at the eye region, monkeys spent significantly less time looking at the eyes in both oxytocin conditions as compared to the saline condition. These results suggest that oxytocin did not necessarily enhance eye looking in capuchins, which is consistent with the results from Pan species, and that endogenous and exogenous oxytocin may behave differently in their effect on how social attention is allocated.

Plasma Testosterone and Androstenedione Levels Follow the Same Sex-Specific Patterns in the Two Pan Species.

Simple SummaryGenerally male mammals are more aggressive than their female peers. In these males, aggressive behavior is linked to levels of androgens; higher levels of testosterone are predictive of higher aggression rates or more severe aggression. There are some species where the pattern of sex-specific aggression is reversed, and it was hypothesized that high levels of androgens may be responsible for social dominance and aggressiveness in these females. Studies so far found that females of species with sex-reversed aggression patterns (e.g., spotted hyenas and ring-tailed lemurs) had lower plasma testosterone levels than their male peers, but a precursor of testosterone, androstenedione, was comparable or even higher in females than in males. This supported the idea that selection for female aggressiveness may be facilitated through augmented androgen secretion. Here we show that in two sister species, bonobos and chimpanzees, that differ in terms of sex-specific aggression patterns, females have lower plasma testosterone levels and higher plasma androstenedione levels than their male peers. Thus, our data do not support a theory of a role of female androgen levels on the expression of sex-specific patterns of aggression.In most animals, males are considered more aggressive, in terms of frequency and intensity of aggressive behaviors, than their female peers. However, in several species this widespread male-biased aggression pattern is either extenuated, absent, or even sex-reversed. Studies investigating potential neuro-physiological mechanisms driving the selection for female aggression in these species have revealed an important, but not exclusive role of androgens in the expression of the observed sex-specific behavioral patterns. Two very closely related mammalian species that markedly differ in the expression and degree of sex-specific aggression are the two Pan species, where the chimpanzee societies are male-dominated while in bonobos sex-biased aggression patterns are alleviated. Using liquid chromatography–mass spectrometry (LC-MS) methods, we measured levels of plasma testosterone and androstenedione levels in male and female zoo-housed bonobos (N = 21; 12 females, 9 males) and chimpanzees (N = 41; 27 females, 14 males). Our results show comparable absolute and relative intersexual patterns of blood androgen levels in both species of Pan. Plasma testosterone levels were higher in males (bonobos: females: average 0.53 ± 0.30 ng/mL; males 6.70 ± 2.93 ng/mL; chimpanzees: females: average 0.40 ± 0.23 ng/mL; males 5.84 ± 3.63 ng/mL) and plasma androstenedione levels were higher in females of either species (bonobos: females: average 1.83 ± 0.87 ng/mL; males 1.13 ± 0.44 ng/mL; chimpanzees: females: average 1.84 ± 0.92 ng/mL; males 1.22 ± 0.55 ng/mL). The latter result speaks against a role of androstenedione in the mediation of heightened female aggression, as had been suggested based on studies in other mammal species where females are dominant and show high levels of female aggressiveness.

Oxytocin promotes species-relevant outgroup attention in bonobos and chimpanzees

Previous research has found that oxytocin (OT) is associated with intergroup behaviour in humans as well as wild chimpanzees, and that exogenous OT affects Pan social attention. The two Pan species, bonobos and chimpanzees, differ drastically from one another in their intensity of intergroup competition, with lethal intergroup aggression often led by males in chimpanzees and more tolerant associations often centered around females in bonobos. However, it remains unclear how exogenous OT changes the two species' responses to ingroup and outgroup individuals. In this study, after intranasal administration of nebulized OT or placebo control, chimpanzees and bonobos viewed image pairs of ingroup and outgroup conspecifics while their eye movements were tracked with an eye-tracker. Although the overall effect of OT was small, we found that OT shifted bonobos' and chimpanzees' attention to outgroup images of the sex primarily involved in intergroup encounters in each species. Specifically, OT selectively shifted attention towards outgroup photos of female conspecifics in bonobos, and those of outgroup male conspecifics in chimpanzees. This suggests that OT generally promotes outgroup attention in both bonobos and chimpanzees but this effect is restricted to the sex most relevant in intergroup relations. These results suggest that, although OT may have a generally conserved role in hominid intergroup behaviour, it may act in species-relevant ways under the influence of their socio-ecological backgrounds.

Drivers of Dyadic Cofeeding Tolerance in Pan: A Composite Measure Approach.

Simple SummaryIn this study on zoo-housed chimpanzees and bonobos, we propose a novel operational measure to quantify how tolerantly animals cofeed around a food source. Using an experimental set-up, we measured five behavioral variables that reflect positive and negative aspects of cofeeding behavior. Using a dimension reduction analysis technique, we found a two-dimensional tolerance model that we labeled “Tolerant Cofeeding” and “Agonistic Cofeeding”. In both species, scores on these two dimensions were predicted by sex, relatedness and social bonds in a similar fashion. High-value relationships between individuals that consist in mutual affiliation and support during aggressive interactions resulted in higher tolerance around food sources, while highly competitive dyads that show more mutual aggression showed lower tolerance. Bonobos and chimpanzees did not differ in their overall cofeeding tolerance, which likely reflects their adaptability to fluctuating social and ecological circumstances. This methodology can now be applied to other animal species to further investigate how variation in social and ecological environments influences the tolerance levels of social bonds, both at the dyadic and group level.This study aimed to construct a composite model of Dyadic Cofeeding Tolerance (DCT) in zoo-housed bonobos and chimpanzees using a validated experimental cofeeding paradigm and to investigate whether components resulting from this model differ between the two species or vary with factors such as sex, age, kinship and social bond strength. Using dimension reduction analysis on five behavioral variables from the experimental paradigm (proximity, aggression, food transfers, negative food behavior, participation), we found a two-factor model: “Tolerant Cofeeding” and “Agonistic Cofeeding”. To investigate the role of social bond quality on DCT components alongside species effects, we constructed and validated a novel relationship quality model for bonobos and chimpanzees combined, resulting in two factors: Relationship Value and Incompatibility. Interestingly, bonobos and chimpanzees did not differ in DCT scores, and sex and kinship effects were identical in both species but biased by avoidance of the resource zone by male–male dyads in bonobos. Social bonds impacted DCT similarly in both species, as dyads with high Relationship Value showed more Tolerant Cofeeding, while dyads with higher Relationship Incompatibility showed more Agonistic Cofeeding. We showed that composite DCT models can be constructed that take into account both negative and positive cofeeding behavior. The resulting DCT scores were predicted by sex, kinship and social bonds in a similar fashion in both Pan species, likely reflecting their adaptability to changing socio-ecological environments. This novel operational measure to quantify cofeeding tolerance can now be applied to a wider range of species in captivity and the wild to see how variation in local socio-ecological circumstances influences fitness interdependence and cofeeding tolerance at the dyadic and group levels. This can ultimately lead to a better understanding of how local environments have shaped the evolution of tolerance in humans and other species.

Do female bonobos (Pan paniscus) disperse at the onset of puberty? Hormonal and behavioral changes related to their dispersal timing

Natal dispersal is a milestone in an animal's life history, but its timing in developmental trajectories may differ between species. Although the two Pan species exhibit a similar pattern of female-biased dispersal, female bonobos (P. paniscus) leave their natal groups at an earlier age than female chimpanzees (P. troglodytes). As a preliminary step to explore the dispersal strategies of female bonobos, this study aimed to determine the relations of sexual swelling development, behavioral and hormonal activation, and first ovulation relative to dispersal timing. We measured levels of urinary estrone conjugates (E1C) and pregnanediol glucuronide (PdG) from 14 nulliparous females in wild bonobo groups at Wamba in the Democratic Republic of the Congo, and recorded their copulations with mature males. When close to dispersal, female bonobos exhibited swelling of the sexual skin (labia minora and perianal region) that did not reach the mature stage. Urinary E1C levels and copulation rates increased slightly before dispersal and greatly increased after dispersal. Ovulatory or gestatory signs implied by daily hormone profiles were not detected until one to two years after dispersal. Our findings indicate that female bonobos disperse at an early pubertal stage before ovulatory cycling is established. This earlier dispersal than sexual maturation could allow female bonobos to postpone reproduction-related energy costs until they become familiar with their new group or gain more time finding the group more suitable for successful reproduction in the future before actually settling. Further demographic and genetic data from dispersal to reproduction will help clarify their dispersal strategies.

Towards the restoration of ancient hominid craniofacial anatomy: Chimpanzee morphology reveals covariation between craniometrics and facial soft tissue thickness.

In modern humans, facial soft tissue thicknesses have been shown to covary with craniometric dimensions. However, to date it has not been confirmed whether these relationships are shared with non-human apes. In this study, we analyze these relationships in chimpanzees (Pan troglodytes) with the aim of producing regression models for approximating facial soft tissue thicknesses in Plio-Pleistocene hominids. Using CT scans of 19 subjects, 637 soft tissue, and 349 craniometric measurements, statistically significant multiple regression models were established for 26 points on the face and head. Examination of regression model validity resulted in minimal differences between observed and predicted soft tissue thickness values. Assessment of interspecies compatibility using a bonobo (Pan paniscus) and modern human subject resulted in minimal differences for the bonobo but large differences for the modern human. These results clearly show that (1) soft tissue thicknesses covary with craniometric dimensions in P. troglodytes, (2) confirms that such covariation is uniformly present in both extant Homo and Pan species, and (3) suggests that chimp-derived regression models have interspecies compatibility with hominids who have similar craniometric dimensions to P. troglodytes. As the craniometric dimensions of early hominids, such as South African australopithecines, are more similar to P. troglodytes than those of H. sapiens, chimpanzee-derived regression models may be used for approximating their craniofacial anatomy. It is hoped that the results of the present study and the reference dataset for facial soft tissue thicknesses of chimpanzees it provides will encourage further research into this topic.

The HLA A03 Supertype and Several Pan Species Major Histocompatibility Complex Class I A Allotypes Share a Preference for Binding Positively Charged Residues in the F Pocket: Implications for Controlling Retroviral Infections.

The major histocompatibility complex (MHC) class I region of humans, chimpanzees (Pan troglodytes), and bonobos (Pan paniscus) is highly similar, and orthologues of HLA-A, -B, and -C are present in both Pan species. Based on functional characteristics, the different HLA-A allotypes are classified into different supertypes. One of them, the HLA A03 supertype, is widely distributed among different human populations. All contemporary known chimpanzee and bonobo MHC class I A allotypes cluster genetically into one of the six HLA-A families, the HLA-A1/A3/A11/A30 family. We report here that the peptide-binding motif of the Patr-A*05:01 allotype, which is commonly present in a cohort of western African chimpanzees, has a strong preference for binding peptides with basic amino acids at the carboxyl terminus. This phenomenon is shared with the family members of the HLA A03 supertype. Based on the chemical similarities in the peptide-binding pocket, we inferred that the preference for binding peptides with basic amino acids at the carboxyl terminus is widely present among the human, chimpanzee, and bonobo MHC-A allotypes. Subsequent in silico peptide-binding predictions illustrated that these allotypes have the capacity to target conserved parts of the proteome of human immunodeficiency virus type 1 (HIV-1) and the simian immunodeficiency virus SIVcpz.IMPORTANCE Most experimentally infected chimpanzees seem to control an HIV-1 infection and are therefore considered to be relatively resistant to developing AIDS. Contemporary free-ranging chimpanzees may carry SIVcpz, and there is evidence for AIDS-like symptoms in these free-ranging animals, whereas SIV infections in bonobos appear to be absent. In humans, the natural control of an HIV-1 infection is strongly associated with the presence of particular HLA class I allotypes. The ancestor of the contemporary living chimpanzees and bonobos survived a selective sweep targeting the MHC class I repertoire. We have put forward a hypothesis that this may have been caused by an ancestral retroviral infection similar to SIVcpz. Characterization of the relevant MHC allotypes may contribute to understanding the shaping of their immune repertoire. The abundant presence of MHC-A allotypes that prefer peptides with basic amino acids at the C termini suggests that these molecules may contribute to the control of retroviral infections in humans, chimpanzees, and bonobos.

Investigating the Function of Mutual Grooming in Captive Bonobos (Pan paniscus) and Chimpanzees (Pan troglodytes)

Social grooming is often exchanged between individuals in many primate species. Rates of bidirectional (or simultaneous mutual) grooming vary across primate species, and its function is not yet fully understood. For example, mutual grooming is frequent in chimpanzees but rare in most primate species including wild bonobos. There are, however, no quantitative data available in captive bonobos. Therefore, through the direct comparison between captive bonobos and chimpanzees, this study aimed to (i) compare the frequency of mutual grooming between Pan species, (ii) explore and compare the function of mutual grooming, and (iii) discuss the rarity of this behavior in wild bonobo populations. We tested three hypotheses following the previous literature in wild chimpanzees. The social bonding hypothesis states that mutual grooming facilitates the maintenance of strong dyadic bonds. The immediate investment hypothesis states that it serves to signal willingness to invest in the interaction. The switching hypothesis states that mutual grooming serves no function but only occurs as an overlap to change the direction of unidirectional grooming. Our findings strongly supported the immediate investment hypothesis, but not the others. Grooming bouts that included mutual grooming were longer and more equitable than bouts without, illustrating that captive Pan species use mutual grooming to maximize their short-term benefits and increase the social value of their interaction. Captive bonobos performed mutual grooming in similar proportions and for similar functions as captive and wild chimpanzees do. This contrasts with wild bonobos who engage in this behavior only rarely. We suggest that the differences in patterns of mutual grooming between captive and wild bonobos might be explained by different degrees of kinship or by a potential intraspecies variation.

Accuracy and spread of nest search behaviour in the Saharan silver ant, Cataglyphis bombycina, and in the salt pan species, Cataglyphis fortis

Desert ants of the genus Cataglyphis are renowned for their navigation abilities, especially for their beeline homing after meandering foraging excursions reaching several hundreds of meters in length. A spiralling nest search is performed when an ant misses the nest entrance upon completing its homebound travel. We examined the nest search behaviours of two desert ant species dwelling in different habitats—Cataglyphis bombycina living in the dunes of the Sahara and Cataglyphis fortis found in the salt pans of North Africa. The two species show distinct differences in walking behaviour. C. bombycina performs a strict tripod gait with pronounced aerial phases, high stride frequencies, and extremely brief ground contact times. In view of these peculiarities and the yielding sand dune substrate, we hypothesised that homing accuracy, and namely distance measurement by stride integration, should be lower in C. bombycina, compared to the well-studied C. fortis with less specialised walking behaviour. We tested this hypothesis in ants’ homebound runs from a feeding site in a linear channel setup. Surprisingly, the accuracies of nest searches were similar in the two ant species, and search accuracy was also independent of the walking substrate, soft dune sand or a hard floor. The spread of the nest search, by contrast, differed significantly between the two species, C. bombycina exhibiting a larger search spread. This may be interpreted as an increased path integration uncertainty due to the above locomotor specialisations, or as a compensation strategy accounting for the silver ants’ particular environmental and behavioural situation.

Why preen others? Predictors of allopreening in parrots and corvids and comparisons to grooming in great apes

AbstractAllogrooming in primates serves not only a hygienic function, but also plays a crucial role in maintaining strong affiliative bonds between group members, which in turn, underpin the emergence of cooperative behavior. In contrast, although allopreening occurs in many avian species, we know little about its social functions. Our study addresses this issue by investigating allopreening in a broad comparative data set including six corvid and nine parrot species. We assessed whether rates of allopreening initiations, proportion of time spent allopreening, and the number of grooming partners in captive group‐housed birds were comparable to patterns observed in captive chimpanzees and bonobos. While parrots and corvids were found to have similar rates of social grooming to bonobos and chimpanzees,Panspecies dedicated significantly more time to social grooming. Animals in larger groups had more grooming partners, but when controlling for the number of potential partners, birds tended to have fewer grooming interaction partners thanPanspecies. We then investigated whether allopreening in parrots and corvids was predicted by behavioral markers of affiliative social bonds (close physical proximity, active feeding, and low levels of agonistic behavior). Results revealed that providing allopreening to a partner was significantly predicted by often being in close proximity, but not engagement in active feeding or agonistic behavior. We examined the region allopreened in a subset of species and found that preening a partner's head was predicted by both close physical proximity and active feeding, while body allopreening was only predicted by close physical proximity. Head preening may confer more hygienic benefits to recipients, and thus, may be more selectively provided to valued partners. Results support the hypothesis that allopreening in corvids and parrots helps maintain social bonds with an individual's most important social partners, showing some similarities to allogrooming in primates.

The position of Neandertal and Homo erectus within the hominid clade based on craniodental morphology and whole mtDNA genomes.

To evaluate the taxonomic position of the Neandertal and Homo erectus within the hominid clade, the variation among and within the hominid taxa was assessed based on the craniodental morphology and integrated with molecular analyses of the whole mtDNA genomes. Ordination and clustering of the Procrustes craniodental landmarks have showed a notable shape transformation from the earliest hominid species to the modern humans. Although levels of distinction between the analyzed taxa (Homo, Pan, Gorilla, and Pongo) are generally corresponding to probable expectations based on their taxonomic rank, few exceptions were found. Notably, the craniodental morphology of Homo erectus showed a greater dissimilarity to other Homo species, where it consistently overlapped or grouped with Pan species on all ordination plots and clustering. In addition, the direct link between European humans and Neandertals, which is well-characterized on all of the phylogenetic trees based on maximum parsimony and maximum likelihood methods, was not outlined in the morphologic-based clustering. Both morphological and molecular distances between Neandertal and modern humans were consistently greater than the distances among modern humans, however, the distances are still smaller than those between any two distinct species (so they are subspecies). The topology of the phylogenetic trees based on the whole mtDNA has shown a minor discrepancy with the results obtained from the craniodental morphologies.

Humans and Chimpanzees Display Opposite Patterns of Diversity in Arylamine N-Acetyltransferase Genes.

Among the many genes involved in the metabolism of therapeutic drugs, human arylamine N-acetyltransferases (NATs) genes have been extensively studied, due to their medical importance both in pharmacogenetics and disease epidemiology. One member of this small gene family, NAT2, is established as the locus of the classic human acetylation polymorphism in drug metabolism. Current hypotheses hold that selective processes favoring haplotypes conferring lower NAT2 activity have been operating in modern humans’ recent history as an adaptation to local chemical and dietary environments. To shed new light on such hypotheses, we investigated the genetic diversity of the three members of the NAT gene family in seven hominid species, including modern humans, Neanderthals and Denisovans. Little polymorphism sharing was found among hominids, yet all species displayed high NAT diversity, but distributed in an opposite fashion in chimpanzees and bonobos (Pan genus) compared to modern humans, with higher diversity in Pan species at NAT1 and lower at NAT2, while the reverse is observed in humans. This pattern was also reflected in the results returned by selective neutrality tests, which suggest, in agreement with the predicted functional impact of mutations detected in non-human primates, stronger directional selection, presumably purifying selection, at NAT1 in modern humans, and at NAT2 in chimpanzees. Overall, the results point to the evolution of divergent functions of these highly homologous genes in the different primate species, possibly related to their specific chemical/dietary environment (exposome) and we hypothesize that this is likely linked to the emergence of controlled fire use in the human lineage.

Metacarpal trabecular bone varies with distinct hand-positions used in hominid locomotion.

Trabecular bone remodels during life in response to loading and thus should, at least in part, reflect potential variation in the magnitude, frequency and direction of joint loading across different hominid species. Here we analyse the trabecular structure across all non-pollical metacarpal distal heads (Mc2-5) in extant great apes, expanding on previous volume of interest and whole-epiphysis analyses that have largely focused on only the first or third metacarpal. Specifically, we employ both a univariate statistical mapping and a multivariate approach to test for both inter-ray and interspecific differences in relative trabecular bone volume fraction (RBV/TV) and degree of anisotropy (DA) in Mc2-5 subchondral trabecular bone. Results demonstrate that whereas DA values only separate Pongo from African apes (Pan troglodytes, Pan paniscus, Gorilla gorilla), RBV/TV distribution varies with the predicted loading of the metacarpophalangeal (McP) joints during locomotor behaviours in each species. Gorilla exhibits a relatively dorsal distribution of RBV/TV consistent with habitual hyper-extension of the McP joints during knuckle-walking, whereas Pongo has a palmar distribution consistent with flexed McP joints used to grasp arboreal substrates. Both Pan species possess a disto-dorsal distribution of RBV/TV, compatible with multiple hand postures associated with a more varied locomotor regime. Further inter-ray comparisons reveal RBV/TV patterns consistent with varied knuckle-walking postures in Pan species in contrast to higher RBV/TV values toward the midline of the hand in Mc2 and Mc5 of Gorilla, consistent with habitual palm-back knuckle-walking. These patterns of trabecular bone distribution and structure reflect different behavioural signals that could be useful for determining the behaviours of fossil hominins.

Alpha and sigma taxonomy of Pan (chimpanzees) and Plio-Pleistocene hominin species

Alpha and sigma taxonomy of Pan (chimpanzees) and Plio-Pleistocene hominin species

Aging and sex affect soluble alpha klotho levels in bonobos and chimpanzees

BackgroundThroughout life, physiological homeostasis is challenged and the capacity to cope with such challenges declines with increasing age. In many species, sex differences exist in life expectancy. Sex-specific differences have been related to extrinsic factors like mate competition and/or intrinsic proximate mechanisms such as hormonal changes. In humans, an intrinsic factor related to aging is soluble alpha klotho (α-Kl). Both sexes show an age-related decline in α-Kl, but throughout life women have higher levels than men of the same age. Sex differences in α-Kl have been linked to a shorter lifespan, as well as to specific morbidity factors such as atherosclerosis and arteries calcifications. In non-human animals, information on α-Kl levels is rare and restricted to experimental work. Our cross-sectional study is the first on α-Kl levels in two long-lived species: bonobos (Pan paniscus) and chimpanzees (Pan troglodytes). As in most mammals, female bonobos and chimpanzees have longer life expectancy than males.MethodsWe measured serum α-Kl levels of 140 subjects from 16 zoos with an ELISA to examine if α-Kl levels reflect this difference in life expectancy.ResultsIn both species and in both sexes, α-Kl levels declined with age suggesting that this marker has potential for aging studies beyond humans. We also found species-specific differences. Adult female bonobos had higher α-Kl levels than males, a difference that corresponds to the pattern found in humans. In chimpanzees, we found the opposite: males had higher α-Kl levels than females.ConclusionWe suggest that contrasting sex differences in adult α-Kl levels mirror the dominance relations between females and males of the two Pan species; and that this might be related to corresponding sex differences in their exposure to stress. In humans, higher cortisol levels were found to be related to lower α-Kl levels. We conclude that there is great potential for studying aging processes in hominoids, and perhaps also in other non-human primates, by measuring α-Kl levels. To better understand the causes for sex differences in this aging marker, consideration of behavioural parameters such as competition and stress exposure will be required as well as other physiological markers.

Food Sharing across Borders

Evolutionary models consider hunting and food sharing to be milestones that paved the way from primate to human societies. Because fossil evidence is scarce, hominoid primates serve as referential models to assess our common ancestors’ capacity in terms of communal use of resources, food sharing, and other forms of cooperation. Whereas chimpanzees form male-male bonds exhibiting resource-defense polygyny with intolerance and aggression toward nonresidents, bonobos form male-female and female-female bonds resulting in relaxed relations with neighboring groups. Here we report the first known case of meat sharing between members of two bonobo communities, revealing a new dimension of social tolerance in this species. This observation testifies to the behavioral plasticity that exists in the two Pan species and contributes to scenarios concerning the traits of the last common ancestor of Pan and Homo. It also contributes to the discussion of physiological triggers of in-group/out-group behavior and allows reconsideration of the emergence of social norms in prehuman societies.

The give and take of food sharing in Sumatran orang-utans, Pongo abelii, and chimpanzees, Pan troglodytes

Proximate factors of primate food sharing, in contrast to its evolutionary explanations, have received little attention. Active food sharing is considered prosocial, since possessors may benefit others by spontaneously passing food or by reacting to ‘signals of need’. However, in contrast to passive sharing, active food sharing is rare in most nonhuman primates. Surprisingly, previous research showed that captive Sumatran orang-utans actively share food more frequently than chimpanzees and bonobos, and hence, appear more prosocial. Yet these comparisons with the two Pan species were relying on previously published studies, which differed with regard to methods and food types used. Here we used the identical procedure and food type to compare the food-sharing behaviour of 10 captive Sumatran orang-utans and 18 chimpanzees, in situations where individuals could monopolize a sharable food source. We focused on communicative behaviours used to initiate food transfers, and assessed whether and how much food was transferred in response to these initiation attempts. In both species, most transfers were initiated by taking the food, resulting in passive sharing, while active sharing by offering food or after requesting it occurred only rarely. However, orang-utans differed from chimpanzees in several aspects. Because the food was mostly monopolized by the adult male, orang-utans attempted to initiate food transfers more frequently, resisted more to taking attempts, and were less likely to transfer whole food items. In both species, requests were less likely to result in food transfers, indicating that in situations involving access to food, they do not necessarily respond to ‘signals of need’. We argue that in addition to instances of active sharing, other factors such as the degree of food monopolization, response rates to ‘signals of need’, and the quality and quantity of transferred food need to be considered to gain a more detailed picture of prosociality across species.