Are bonobos (Pan paniscus) really more bipedal than chimpanzees (Pan troglodytes)?

Of the living apes, the chimpanzee (Pan troglodytes) and bonobo (Pan paniscus) are often presented as possible models for the evolution of hominid bipedalism. Bipedality in matched pairs of captive bonobos and chimpanzees was analyzed to test hypotheses for the evolution of bipedalism, derived from a direct referential model. There was no overall species difference in rates of bipedal positional behavior, either postural or locomotory. The hominoid species differed in the function or use of bipedality, with bonobos showing more bipedality for carrying and vigilance, and chimpanzees showing more bipedality for display.     

Sex Differences in Object Manipulation in Wild Immature Chimpanzees (Pan troglodytes schweinfurthii) and Bonobos (Pan paniscus): Preparation for Tool Use?

Sex differences in immatures predict behavioural differences in adulthood in many mammal species. Because most studies have focused on sex differences in social interactions, little is known about possible sex differences in ‘preparation’ for adult life with regards to tool use skills. We investigated sex and age differences in object manipulation in immature apes. Chimpanzees use a variety of tools across numerous contexts, whereas bonobos use few tools and none in foraging. In both species, a female bias in adult tool use has been reported. We studied object manipulation in immature chimpanzees at Kalinzu (Uganda) and bonobos at Wamba (Democratic Republic of Congo). We tested predictions of the ‘preparation for tool use’ hypothesis. We confirmed that chimpanzees showed higher rates and more diverse types of object manipulation than bonobos. Against expectation, male chimpanzees showed higher object manipulation rates than females, whereas in bonobos no sex difference was found. However, object manipulation by male chimpanzees was play-dominated, whereas manipulation types of female chimpanzees were more diverse (e.g., bite, break, carry). Manipulation by young immatures of both species was similarly dominated by play, but only in chimpanzees did it become more diverse with age. Moreover, in chimpanzees, object types became more tool-like (i.e., sticks) with age, further suggesting preparation for tool use in adulthood. The male bias in object manipulation in immature chimpanzees, along with the late onset of tool-like object manipulation, indicates that not all (early) object manipulation (i.e., object play) in immatures prepares for subsistence tool use. Instead, given the similarity with gender differences in human children, object play may also function in motor skill practice for male-specific behaviours (e.g., dominance displays). In conclusion, even though immature behaviours almost certainly reflect preparation for adult roles, more detailed future work is needed to disentangle possible functions of object manipulation during development.     

How Precisely Do Bonobos (Pan paniscus) Grasp Small Objects?

The general objective of this study was to compare the precise grasping behavior and intermanual differences in performance between three Pan paniscus and five Homo sapiens in grasping small objects. We compared the temporal pattern of two submovements of consecutive grasping cycles, the (visuomotor) reaching and the (sensorimotor) grasping. Both species were similarly successful in this task, they showed a behavioral right-hand preference and preferred specific types of grips. Bonobos required less time for reaching an object but a much longer time to grasp it than humans did. Thus, the species pursued different strategies. We assumed that this might be due to the different grip techniques. However, grip preferences did not serve a quicker intramanual performance but they pronounced differences between hands. Intermanual differences in timing were restricted to the reaching part and more strongly in bonobos than in humans. However, the right hand need not necessarily perform quicker. As in the case of humans, we assume that attentional cues were focused more on preparing a proper grip with the right hand than on a quick performance. However, strong intermanual differences in bonobos may indicate an overall stronger neuronal asymmetry in the motor organization of the finger musculature that prepare a proper grip than is true of humans.     

Chimpanzees’ constructional praxis (Pan paniscus, P. troglodytes)

This study investigated chimpanzees spontaneous spatial constructions with objects and especially their ability to repeat inter-object spatial relations, which is basic to understanding spatial relations at a higher level than perception or recognition. Subjects were six chimpanzees—four chimpanzees and two bonobos—aged 6–21 years, all raised in a human environment from an early age. Only minor species differences, but considerable individual differences were found. The effect of different object samples was assessed through a comparison with a previous study. A common overall chimpanzee pattern was also found. Chimpanzees repeated different types of inter-object spatial relations such as insertion (I), or vertical (V), or next-to (H) relations. However chimpanzees repeated I or V relations with more advanced procedures than when repeating H relations. Moreover, chimpanzees never repeated combined HV relations. Compared with children, chimpanzees showed a specific difficulty in repeating H relations. Repeating H relations is crucial for representing and understanding multiple reciprocal spatial relations between detached elements and for coordinating independent positions in space. Therefore, the chimpanzees difficulty indicates a fundamental difference in constructive space in comparison to humans. The findings are discussed in relation to issues of spatial cognition and tool use.     

Male–Female Association Patterns Among Free-ranging Chimpanzees (Pan troglodytes schweinfurthii)

Although male–female relationships can offer a number of advantages such as protection or social support, they are poorly studied among primates compared to same-sex relationships. We used 12 yr of data from the Kanyawara chimpanzee community to compare three independent measures of association (party association, 5m association, and grooming) among all adult dyads. Party association exhibited by male–female dyads was of intermediate strength between strong male–male and weak female–female association. Male–female dyads were less likely to be within 5m of one another and to groom as male–male dyads, but equally likely to be within 5m and more likely to groom as female–female dyads. Variation in male–female association strength was not related to male rank but was affected by female ranging patterns and female reproductive states. Females with core areas in the center of the home range were more likely to be in parties with males but did not show higher spatial proximity or grooming indices compared to females ranging in the periphery. Party association and 5m indices were higher for dyads of males and estrous females compared to those with anestrous females. These results indicate that male–female dyads are likely to associate with one another more often than female–female dyads because of overlapping ranging patterns and short-term changes in female reproductive state. We conclude that male and female chimpanzees do not exhibit proximity and grooming patterns indicative of strong affiliative bonds. This study also highlights the importance of using multiple independent measures of bond strength in studies of primate social dynamics.     

RNA decay machines: Deadenylation by the Ccr4–Not and Pan2–Pan3 complexes

Shortening and removal of the 3′ poly(A) tail of mature mRNA by poly(A)-specific 3′ exonucleases (deadenylases) is the initial and often rate-limiting step in mRNA degradation. The majority of cytoplasmic deadenylase activity is associated with the Ccr4–Not and Pan2–Pan3 complexes. Two distinct catalytic subunits, Caf1/Pop2 and Ccr4, are associated with the Ccr4–Not complex, whereas the Pan2 enzymatic subunit forms a stable complex with Pan3. In this review, we discuss the composition and activity of these two deadenylases. In addition, we comment on generic and specific mechanisms of recruitment of Ccr4–Not and Pan2–Pan3 to mRNAs. Finally, we discuss specialised and redundant functions of the deadenylases and review the importance of Ccr4–Not subunits in the regulation of physiological processes. This article is part of a Special Issue entitled: RNA Decay mechanisms.     

Does space in the jaw influence the timing of molar crown initiation? A model using baboons (Papio anubis) and great apes (Pan troglodytes, Pan paniscus)

Radiographic and histological studies of baboon (Papio hamadryas, P. anubis) and chimpanzee (Pan troglodytes) permanent tooth development have found that periods of molar crown mineralization overlap markedly in chimpanzees but are staggered in baboons. Here we test the hypothesis that these intertaxon differences in molar initiation are primarily due to the space available in the mandibles of each species for these teeth. This study includes radiographic, linear measurement, and three-dimensional (3D) coordinate landmark data taken from baboon (Papio anubis n=51) and great ape (Pan paniscus n=43, P. troglodytes n=60) mandibles and permanent molars across a broad developmental range for each taxon. Unexpectedly, 3D multivariate statistical shape analysis of the molar crypt, crown, and root data shows that all three species trajectories of molar row shape change are indistinguishable from each other. Qualitative analysis of these 3D data reveals subtle and inconclusive intergeneric differences in the space maintained between adjacent molars during growth. The space distal to each newly initiated molar is slightly greater in the baboon. Bivariate analyses comparing molar row and mandibular corpus proportions in Papio and Pan fail to show clear or consistent taxonomic differences in the ratio of space afforded developing molars in the alveolar bone. Thus, there is a poor correlation between mandibular proportion and both intermolar spacing and 3D molar development pattern. Contrary to earlier studies, these results suggest that pattern of molar crown initiation and temporal overlap of adjacent mineralizing crowns is not significantly different between Papio and Pan. This may be due in part to the inclusion here of not only 3D molar crown data but also 3D molar crypt data. This study strongly refutes the hypothesis that space available in the mandible directly underlies different times of permanent molar crown initiation between Papio and Pan.     

Genetic structure of a rare European conifer, Serbian spruce (Picea omorika (Panč.) Purk.)

Genetic variation in 13 populations of a Balkan endemic, Serbian spruce (Picea omorika (Panč.) Purk.), was investigated using 16 isozyme loci. Serbian spruce is characterized by low levels of genetic variation (average proportion of polymorphic loci was 20.9% and average expected heterozygosity was 0.067). In most populations, a significant surplus of heterozygotes was observed, indicating a strong selection against inbreds. In the largest populations, fixation indices were positive, probably due to within-population differentiation and the associated Wahlund effect. Despite a strong overall differentiation (F_ST=0.261), no geographical trends in the genetic variation could be identified. Genetic drift caused by small effective population sizes and a strong fragmentation is the most plausible explanation for such variation patterns. A Bayesian analysis of population structure revealed the existence of two clusters, which are supposed to be possible remnants of ancient differentiation within a large range of the predecessor of Serbian spruce, Picea omoricoides Weber. Ex situ conservation measures, namely establishing seed orchards from trees of mixed origin, are proposed as a necessary complement to in situ protection of natural stands.     

The sensitivity of embryogenic tissue of Picea omorika (Panč.) Purk. to antibiotics

In an attempt to develop a transformation procedure for somatic embryos of Picea omorika (Pan.) Purk. using Agrobacterium, the sensitivity of non-transformed embryogenic tissues of P. omorika to antibiotics was measured. Two groups of antibiotics were tested: antibiotics commonly used to eliminate Agrobacterium from tissue culture (carbenicillin and cefotaxime), and antibiotics for the selection of transformed tissue (kanamycin, paromomycin, neomycin and hygromycin). Carbenicillin (500 mg l^–1) reduced proliferation of P. omorika embryogenic tissue by 50% as compared to the control. Cefotaxime (500 mg l^–1) was less toxic to embryogenic tissue growth (93% of the control) and is thus more suitable for the elimination of A. tumefaciens from embryogenic tissue of P. omorika. Embryogenic tissue showed severe susceptibility to kanamycin and hygromycin. Induction of secondary embryogenesis was blocked by 10 mg l^–1 of kanamycin or 2 mg l^–1 of hygromycin. The aminoglycoside analogs of kanamycin, paromomycin and neomycin inhibited embryogenic tissue induction at concentrations of 5 and 100 mg l^–1, respectively. The higher tolerance of embryogenic tissue to neomycin indicated that of the antibiotics tested neomycin may be most useful for selection of nptII-transformed embryogenic tissue of Picea omorika.     

Versuche zur Symbol-Ereignis-Verknüpfung bei einer Zwergschimpansen (Pan paniscus Schwarz, 1929)

According toFiedler (1956) the pygmy chimpanzee is considered to be a separate species of the pongidae. Therefore, our investigations with a young female pygmy chimpanzee had two special aims. First, we hoped to collect some facts concerning the transferability of the experiments ofPremack (1971) within the great apes; he investigated the language skills of a common chimpanzee. Besides, we wanted to test the intellectual abilities of this littel known primate species. Pieces of wood, differing in color and shape, were used as symbols. At both sides, there was a magnet to hang them on an iron board. The subject had to choose between two or more signs out of three groups: food symbols, activity symbols and quantity symbols. The hanging up of one, later of two correlated symbols was followed by a special event for each of them. There is high concordance between the subject’s preferring of events and its preferring of symbols. The correlation-coefficient for food and food signs is as high as 0.95. Nearly without mistake is the subject’s use of the much/little signs in connection with the food symbols. The rate of error in general is between 10–20% and is in accordance to the corresponding results mentioned byPremack (1971). The way the subject learned the meaning of the symbols changed from slow to sudden comprehension. In general, these results provide further evidence of the high intelligence level of pygmy chimpanzees.      

Tool-use by chimpanzees (Pan t. troglodytes) in the Lopé Reserve,Gabon

During a long-term study of chimpanzees in the Lopé Reserve, Gabon, tool-use to obtain insects or their products was observed four times and on 23 other occasions tools made and used by chimpanzees were recovered. Of 144 tools used to obtain insects, 142 were made from woody material. Tools varied in dimensions and in the degree of modification (bark stripping and fraying of ends) but those used to obtain different prey species were generally similar and resembled tools described from other chimpanzee populations in central Africa. Use of a leaf “sponge” to obtain water from a hole in the branch of a tree was observed once. Four tool-use patterns shown by other chimpanzee populations appeared to be absent at Lopé as no direct or indirect evidence of their existence was obtained in ten years.     

Spatiotemporal Variation in Bonobo (Pan paniscus) Habitat Use in a Forest–Savanna Mosaic

International Journal of Primatology - Understanding the ecological and behavioral variation of primates is central to improving conservation strategies. Studies of chimpanzees (Pan troglodytes)...     

Survey of savanna chimpanzees (Pan troglodytes verus) in Southeastern Sénégal

A survey of the western subspecies of chimpanzee (Pan troglodytes verus) was conducted from 1 February to 9 April 2000 in Sénégal, West Africa, by the Miami Assirik Pan Project (MAPP). In addition to the Assirik area of the Parc National du Niokolo Koba (PNNK), areas south and east of the park were surveyed. Nests made by chimpanzees were used to estimate chimpanzee distribution and densities. Within the PNNK, chimpanzees were estimated to occur at an average of 0.13 individuals/km(2). Chimpanzee nests were recorded in nine of 10 locales surveyed outside of the PNNK, as well as within the park. Data on 994 nests made by chimpanzees were recorded outside the PNNK, while 736 nests were recorded inside the park. Nest density in areas surveyed outside the PNNK, such as Bandafassi, Tomboronkoto, and Segou, was comparable to that of Assirik in habitats where nests were concentrated (i.e., evergreen gallery forest). The purpose of MAPP was to initiate long-term research of chimpanzees in southeastern Sénégal, as a follow-up to the Stirling African Primate Project (SAPP) of the 1970s. We sought to replicate the standards set by the SAPP project, except when technological innovations allowed improvement in data collection procedures (e.g., the global positioning system (GPS)).     

Benzyladenine induction of buds and somatic embryogenesis in Picea omorika (Pančić) Purk.

Somatic embryogenesis and adventitious bud formation, initiated from shoot explants of Picea omorika is described. Benzyladenine (BA) as the only growth regulator, added to modified Von Arnold and Eriksson medium, induced formation of both adventitious buds and embryogenic tissue. Optimal BA concentration for bud induction was 4.5 M and further bud development and plantlet formation was achieved on growth regulator-free medium. The embryogenic tissue formation was induced when the explants were first grown on the medium with high BA content (22.5 M) and then transferred to medium without growth regulators. Subsequent proliferation of embryogenic tissue was accomplished by subculturing on medium containing 9 M 2,4-dichlorophenoxyacetic acid and 4.5 M BA, and further embryo development was achieved on medium with 12 M abscisic acid. Embryos cultured on growth regulator-free medium formed roots and rooted plantlets were successfully established in soil in the greenhouse.     

Chaînes opératoires and resource-exploitation strategies in chimpanzee (Pan troglodytes) nut cracking

We apply archaeological methods to extend our knowledge of chimpanzee material culture. The cha?ne opératoire conceptual framework, as introduced by ethnography, established technology as a phased process. Prehistoric archaeology adopted this concept to elucidate technological variability in tool-making procedures, based on knowledge of tool functions or subsistence patterns. We focused on the detection of operational sequences by wild chimpanzees (Pan troglodytes verus) when nut cracking with lithic implements at the sites of Bossou and Diecké, Guinea, West Africa. Thus, while it has recently been claimed that chimpanzees leave behind recognizable assemblages of stone hammers that can be morphologically distinguished from Oldowan hammers, this is the first study to focus specifically on the existence of operational sequences during the utilization of stone tools by wild chimpanzees. By combining primatological and archaeological methods and examining ecological areas inhabited by different chimpanzee groups, we sought technological variability and identified variables influencing regional diversity in tool typology and technology. We compared three case studies: (1) Bossou-direct recording of experimental nut-cracking sessions; (2) Bossou- direct and indirect monitoring of nut-cracking sites in the wild; (3) Diecké-indirect monitoring of nut-cracking sites in the wild. Results suggest that chimpanzees perform sequences of repeated tool transport and nut cracking. Data show discrimination of tool functions based on tool features. We identified the most technologically complex tool for nut cracking, which was composed of four stones. We found regional diversity in chimpanzee stone assemblages. Raw-material type and tool mobility constrain technological development in human and nonhuman primates. Spatial analysis of tool distribution indicates a pattern of resource-exploitation strategy, revealing affinities with Oldowan.     

Fruit sharing between wild adult chimpanzees (Pan troglodytes schweinfurthii): a socially significant event?

Although food sharing is a habitual aspect of chimpanzee (Pan troglodytes) life, sharing of plant foods between unrelated adults is rare. Observations of such behavior have typically been interpreted as the outcome of a process by which individuals that are otherwise unable to gain access to the food manage to obtain a nutritional benefit. Here we present behavioral details and an acoustic analysis regarding an observation of food sharing between unrelated adult wild chimpanzees that we suggest cannot be explained using traditional nutrition-based models. Instead we propose that the exchange is only understandable as a socially important event, and we cite two further observations in the same population that support this suggestion.     

How did bonobos come to range south of the congo river? Reconsideration of the divergence of Pan paniscus from other Pan populations

While investigating the genetic structure in wild bonobos,¹ we realized that the widely accepted scenario positing that the Pleistocene appearance of the Congo River separated the common ancestor of chimpanzees (Pan troglodytes) and bonobos (P. paniscus) into two species is not supported by recent geographical knowledge about the formation of the Congo River. We explored the origin of bonobos using a broader biogeographical perspective by examining local faunas in the central African region. The submarine Congo River sediments and paleotopography of central Africa show that the Congo River has functioned as a geographical barrier for the last 34 million years. This evidence allows us to hypothesize that when the river was first formed, the ancestor of bonobos did not inhabit the current range of the species on the left bank of the Congo River but that, during rare times when the Congo River discharge decreased during the Pleistocene, one or more founder populations of ancestral Pan paniscus crossed the river to its left bank. The proposed scenario for formation of the Congo River and the corridor hypothesis for an ancestral bonobo population is key to understanding the distribution of great apes and their evolution.     

Behavioural responses to photographs by pictorially naïve baboons (Papio anubis), gorillas (Gorilla gorilla) and chimpanzees (Pan troglodytes)

This study assessed how pictorially naïve nonhuman primates understand pictures. Fifty-five baboons with no prior exposure to pictures were trained to grasp a slice of banana presented against a pebble in a two alternative forced choice task. Post-training testing involved three stimulus pairs: (1) real banana slice vs. its picture, (2) the banana picture vs. a real pebble and (3) banana picture vs. a pebble picture which were presented twice. Preliminary data were also collected on naïve gorillas (n = 4) and chimpanzees (n = 7) using the same procedure. Baboons revealed a preference for the food picture in (2) and (3) and often ate this stimulus, but the food item and its picture were accurately discriminated in (1). These results suggest that baboons mistook the pictorial stimulus and its referent, but processed the banana pictures as poor exemplars of the real banana category. Among apes, only gorillas ate the banana pictures, suggesting that picture–object confusion may also occur in this species. Findings are discussed as pertaining to the general issue of representational abilities in nonhuman primates, and its evolution.