Nut cracking by wild chimpanzees in Sierra Leone,West Africa

I report an incidence of chimpanzee (Pan troglodytes verus) cracking ofDetarium senegalense (Caesalpiniaceae) nuts on Tiwai Island, Sierra Leone. Similar behavior has been reported from other West African locations, but not from Central or East Africa.     

Cranial capacity estimations of the FrankfurtPan troglodytes verus collection

Cranial capacity measurements of 18 individuals (9 ♂, 9 ♀) of a total collection of 277Pan troglodytes verus skulls were taken using four different methods:- two conventional filling methods and two mathematical methods. The two mathematical analyses normally used in hominids, were for the first time specifically modified forPan. The results probably present a more accurate estimation of total cranial capacity inPan and specifically inPan troglodytes verus (352 cm³) than previously available. Sexual differences related to this trait were also measured. Regardles of the methods used,Pan troglodytes verus seems to manifest the smallest cranial capacity of all subspecies ofPan.     

Growth of wild and laboratory born chimpanzees

The skeletal remains of a wild juvenile chimpanzee,Pan troglodytes verus, of known chronological age are measured and found to be smaller than laboratory born and fed juveniles of the same age. Other wild born immature skeletal materials of all the three subspecies ofPan troglodytes, including both known and estimated chronological ages, are also smaller than laboratory born chimpanzees when comparisons are made on corresponding age groups. Differences between wild and laboratory born chimpanzees are larger in the limb bones than in the cranium. Limb bones of laboratory individuals grow earlier than those of wild ones regardless of subspecies. Small limb bone size of wild chimpanzees is discussed in terms of life processes.     

Laparoscopic repair of an umbilical hernia in a Western chimpanzee (Pan troglodytes verus) rescued in Sierra Leone

Laparoscopic umbilical hernia repair was performed in an adult female Western chimpanzee (Pan troglodytes verus) in a rescue centre in Sierra Leone. Biological mesh and laparoscopic spiral tacks via three 5 mm laparoscopy ports were used to close a 4 cm hernia. The chimpanzee recovered quickly and the hernia did not recur.     

Locking plate femur fracture repair in a juvenile Western chimpanzee (Pan troglodytes verus) in Sierra Leone

This case reports the successful management of a traumatic diaphyseal femoral fracture in an infant Western chimpanzee (Pan troglodytes verus) from a rescue centre in Sierra Leone using a 2.4-mm intramedullary pin and two 2.7-mm String of Pearls^™ locking plates. Locking plate use has not been previously described in chimpanzees.     

Case report on the death of a wild chimpanzee (Pan troglodytes verus)

The present paper gives a case report on the death of a wild young chimpanzee (Pan troglodytes verus) at Bossou, Guinea. The corpse of a 6.5-year-old male was found, and an autopsy suggested that he had died from some non-epidemic disease or from poisoning. Morphological measurements of the skeleton revealed that the chimpanzee was much smaller than corresponding individuals in captivity. The dental formula of the chimpanzee coincided with those of 5- to 5.5-year-olds in captivity. The death of this chimpanzee suggests that some of the individuals who had disappeared at Bossou had died of natural causes.     

Technical note: Chromosomal and mtDNA analysis of Oliver

Oliver is an African ape whose species identity has been debated in the popular media and by various scientists since the early 1970s. Although decisive morphological data has never been adduced on Oliver, many reports indicated that Oliver was morphologically unusual for a chimpanzee, particularly in his habitual bipedal posture. In addition, his diploid chromosome number was reported to be inconsistent with either human or chimpanzee, but instead intermediate between those species. We performed standard chromosomal studies which demonstrated that Oliver had the diploid number expected for a chimpanzee (2N = 48) and that the banding patterns of his chromosomes were typical for a chimpanzee and different from both humans and bonobos. We also sequenced a 312 bp region of his mitochondrial DNA D-loop region. Results indicated a high sequence homology to the Central African variety of chimpanzee, Pan troglodytes troglodytes. The highest percent homology was observed with a previously characterized specimen from Gabon, strongly suggesting that Oliver originated from this region. Am J Phys Anthropol 105:395–403, 1998. © 1998 Wiley-Liss, Inc.     

Nasal region polymorphism frequencies in the FrankfurtPan troglodytes verus collection

The results of a morphometric study on the mid-facial region in a collection of 278Pan troglodytes verus skulls are reported. Upwardly divergent nasal bones were found in 35 of the 124 specimens (28.2%) in which this feature could be analyzed. Appreciable separation (1 cm. or more) between nasion and glabella characterized 13 of the 154 skulls (8.4%) in which both these osteometric points could be observed. These findings document the wide ranges of variation that are normally to be found in modest-sized population samples of hominoid primates, and point to the inherent unreliability of cladistic analyses based on the hypothesis that hominoid primate taxa are monomorphic for these character states in the nasal region.     

Morphological variation in adult chimpanzees (Pan troglodytes verus) of the Taï National Park, Côte d'Ivoire

Twenty five adult chimpanzee skeletons (Pan troglodytes verus) of known age and sex (15 females, 10 males) from a long‐term study site in Taï National Park, Cote d'Ivoire present new data on variation. These skeletons provide a rare opportunity to measure the cranium and postcranium from the same individuals. We compare measurements and indices of the Taï sample with those of relatively complete Pan troglodytes schweinfurthii skeletons from Gombe National Park, Tanzania. Measurements of Pan paniscus are included as an outside comparison. The Taï and Gombe samples are analyzed by sex; combined sex samples are compared between the two groups, and the two sexes to each other. Taï females and males do not differ in most long bone lengths or in pelvic dimensions, but do differ significantly in cranial capacity, facial measurements, clavicle length, scapular breadth, and femur length. Gombe females and males differ significantly in some facial measurements and in scapular breadth. In combined sex samples, Taï individuals have lower cranial capacity, longer palate and mandible, and greater dimensions in the trunk and limb lengths. Taï females account for most of the variation; males differ from each other only in greater length of humerus and femur. The Taï skeletons provide new data for assessing individual variation and sexual dimorphism within and between populations and species. The combination of cranial and postcranial data provides a clearer picture of chimpanzee intraspecific and interspecific variation than can be gained from either data set alone. Am J Phys Anthropol, 2008. © 2007 Wiley‐Liss, Inc.     

Enamel hypoplasias as indicators of developmental stress in pongids and hominids

This paper reports the results from a study of enamel hypoplasia in the deciduous dentition of free-living Liberian chimpanzees (Pan troglodytes verus). The entire study collection includes 280 specimens (278 skulls plus two unassociated mandibles), of which 70 are young enough to retain some decidous teeth. Among the subsample of infants, the total frequency of hypoplasia summed over all teeth in any individual reaches 80%, being expressed in the form of pits rather than transverse lines. Hypoplasia of a linear form was common in the permanent dentition, attaining a frequency of 46.7% on maxillary central incisors and 69.7% on mandibular canines.     

A longitudinal study of urinary dipstick parameters in wild chimpanzees (Pan troglodytes verus) in Côte d'Ivoire

We performed 796 dip‐stick tests on urine from 100 wild West African chimpanzees (Pan troglodytes verus) from 4 habituated groups in the tropical rain forest of Taï National Park, Cote d'Ivoire, to establish reference values for health monitoring. Specific gravity was also measured on 359 urine samples from 62 chimpanzees. The effect of age, sex, group, month, estrus, pregnancy, meat consumption, and acute respiratory disease on pH, leucocytes, protein, blood, hemoglobin, and glucose was examined using ordinal logistic regression. The presence of nitrite, ketones, bilirubin, and urobilinogen in urine was also recorded. Outbreak of acute respiratory disease did not influence any of the urinary parameters. Thirty‐seven percent of the samples had a pH <7 and the whole range of pH was found through the year, in all age groups, and in both sexes. Meat consumption lowered the urinary pH. Our results show that all pH levels must be considered normal for the West African chimpanzee subspecies P. troglodytes verus living in the rainforest. We also found a cluster of glucose‐positive samples at a specific point in time which was not attributed to diabetes mellitus. These findings highlight that there are differences in normal physiological parameters among wild chimpanzees living in different habitats. Am. J. Primatol. 72:689–698, 2010. © 2010 Wiley‐Liss, Inc.     

Oil Palm Use by Adjacent Communities of Chimpanzees at Bossou and Nimba Mountains,West Africa

We investigated oil palm (Elaeis guineensis) use for feeding in 3 chimpanzee communities: Bossou and Seringbara in Guinea and Yealé in Côte d'Ivoire. Bossou was used as the benchmark for comparison. Bossou chimpanzees (Pan troglodytes verus) exhibit a wide range of oil palm targeted behaviors. We used direct observations of their two tool use, i.e., nut-cracking and pestle pounding, to establish strict and reliable criteria to ascertain the presence of comparable behaviors at the two adjacent Nimba sites. Based on monthly surveys of oil palms across the three sites, significant differences in patterns of use emerged. Bossou chimpanzees demonstrated the greatest frequency of oil palm use, while Seringbara chimpanzees, 6 km away, failed to exhibit any use and Yealé chimpanzees, 12 km away, showed all uses comparable to Bossou chimpanzees except pestle pounding and mature leaf pith-feeding. We examined the density and distribution of oil palms, tool availability for nut-cracking and pestle pounding, fruit, flower and nut availability, competition with sympatric species for fruit and nuts and the diversity of fruit species in the diet across the 3 sites. We found no clear difference in proximate environmental variables underlying observed variations in oil palm use among the 3 sites, yielding the conclusion that the differences are cultural. Assuming individual interchange between communities and the involvement of social learning in the intracommunity transmission and maintenance of oil palm uses, the result raises interesting questions about diffusion of behavior between neighboring chimpanzee communities.     

Dental caries in wild primates: Interproximal cavities on anterior teeth

Dental caries has been reported in a variety of primates, although it is still considered rare in wild populations. In this study, 11 catarrhine primate taxa (n = 339 individuals; 7946 teeth) were studied for the presence of caries. A differential diagnosis of lesions in interproximal regions of anterior teeth was undertaken, since they had been previously described as both carious and non-carious in origin. Each permanent tooth was examined macroscopically, with severity and position of lesions recorded. Two specimens were examined further, using micro-CT scans to assess demineralization. Differential diagnosis confirmed the cariogenic nature of interproximal cavities on anterior teeth (ICATs). Overall results show 3.3% of all teeth (i.e., anterior and posterior teeth combined) were carious (n = 262), with prevalence varying among species from 0% to >7% of teeth affected. Those with the highest prevalence of ICATs include Pan troglodytes verus (9.8% of anterior teeth), Gorilla gorilla gorilla (2.6%), Cercopithecus denti (22.4%), Presbytis femoralis (19.5%), and Cercopithecus mitis (18.3%). ICATs make up 87.9% of carious lesions on anterior teeth. These results likely reflect dietary and food processing differences among species, but also between the sexes (e.g., 9.3% of all female P. troglodytes verus teeth were carious vs. 1.8% in males). Processing cariogenic fruits and seeds with the anterior dentition (e.g., wadging) likely contributes to ICAT formation. Further research is needed in living primate populations to ascertain behavioral/dietary influences on caries occurrence. Given the presence of ICATs in frugivorous primates, their diagnosis in archaeological and paleontological specimens may shed light on diet and food processing behaviors in fossil primates.     

Grouping Patterns and Competition Among Female <Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis> at Ngogo,Kibale National Park,Uganda

In mammals, access to mates is probably the most important influence on male reproductive success, whereas foraging efficiency is probably the most important influence on female reproductive success Emlen and Oring (Science 197:215–223, 1977). Male chimpanzees (Pan troglodytes) are highly gregarious and form cooperative relationships with other males. In contrast, female social relationships vary within and between populations. Females in most East African populations, e.g., Gombe, Mahale, Kibale-Kanyawara, are less gregarious than males and spend most of their time alone or with only their dependent offspring. Researchers have attributed low female gregariousness to the high potential for feeding competition. I provide the first data on association patterns and agonistic interactions of female chimpanzees (Pan troglodytes schweinfurthii) from the unusually large Ngogo community, Kibale National Park, Uganda. Ngogo females were less gregarious than males, but spent a mean of 64% of their time in association with ≥1 other females and as much time in all-female parties as they did alone. Further, female dyads associated nonrandomly and they formed associative cliques. Association levels within cliques were similar to those among the relatively gregarious West African chimpanzee females at Taï (Pan troglodytes verus) and among bonobo (P. paniscus) females. Agonistic interfemale interactions were extremely rare, and monthly mean party size and the numbers of anestrous females per party do not correlate significantly with fruit availability. Thus, Ngogo females maintained relatively high levels of gregariousness, but avoided detrimental feeding competition by preferentially associating with a small subset of other community females.     

Unconstrained cranial evolution in Neandertals and modern humans compared to common chimpanzees

A variety of lines of evidence support the idea that neutral evolutionary processes (genetic drift, mutation) have been important in generating cranial differences between Neandertals and modern humans. But how do Neandertals and modern humans compare with other species? And how do these comparisons illuminate the evolutionary processes underlying cranial diversification? To address these questions, we used 27 standard cranial measurements collected on 2524 recent modern humans, 20 Neandertals and 237 common chimpanzees to estimate split times between Neandertals and modern humans, and between Pan troglodytes verus and two other subspecies of common chimpanzee. Consistent with a neutral divergence, the Neandertal versus modern human split-time estimates based on cranial measurements are similar to those based on DNA sequences. By contrast, the common chimpanzee cranial estimates are much lower than DNA-sequence estimates. Apparently, cranial evolution has been unconstrained in Neandertals and modern humans compared with common chimpanzees. Based on these and additional analyses, it appears that cranial differentiation in common chimpanzees has been restricted by stabilizing natural selection. Alternatively, this restriction could be due to genetic and/or developmental constraints on the amount of within-group variance (relative to effective population size) available for genetic drift to act on.     

On the nomenclature ofPan paniscus

In 1929 a subspecies of chimpanzee was classified asPan satyrus paniscus, a subspecies of theeastern chimpanzee, and elevated to species level,Pan paniscus, in 1933. Review of the literature indicates that this ape type was known since 1881 from several locations throughout the Congo Basin and first scientifically described in 1887 asTroglodytes niger var.marungensis, a subspecies of thewestern chimpanzee. The evidence presented here demonstrates that the synonymmarungensis was known and used as an earlier name. According to the International Code of Zoological Nomenclature, the earliest name awarded to a species is the one which has to be recognized in use. However, maintaining the emphasis on stability and assuming that to change the name would cause considerable confusion in the literature, it is this author's recommendation that the earlier namemarungensis be suppressed and the later synonympaniscus be conserved while an application is being considered for ruling by the International Commission on Zoological Nomenclature.     

A complete sequence of the mitochondrial genome of the western lowland gorilla

The complete mitochondrial DNA (mtDNA) molecule of the gorilla was sequenced. The entire sequence, 16,412 nucleotides, was determined by analysis of natural (not polymerase chain reaction) restriction fragments covering the whole molecule. The sequence was established from one individual and thus nonchimeric. After comparison with the COII gene of gorilla specimens with known geographical origin, the sequence was identified as characteristic of the Western lowland gorilla, Gorilla gorilla gorilla. With the exception of the NADH2 gene, all genes have a methionine start codon. The inferred start codon of NADH2 is ATT (isoleucine). The COIII, NASDH4, and cytochrome b genes are not terminated by a stop codon triplet, and the COI gene is probably terminated by an AAA triplet rather than by a regular stop codon. The great majority of genic sequences (rRNAs, peptide-coding genes, tRNAs) of the complete mtDNAs of Gorilla, Pan, and Homo show a greater similarity between Pan and Homo than between either of these genera to Gorilla. The analysis of the peptide-coding genes suggest that relative to comparison between Homo and Pan a certain degree of transition saturation has taken place in codon position 3 in comparisons between Gorilla to either Homo or Pan.      

Chimpanzee fetal G gamma and A gamma globin gene nucleotide sequences provide further evidence of gene conversions in hominine evolution

The fetal globin genes G gamma and A gamma from one chromosome of a chimpanzee (Pan troglodytes) were sequenced and found to be closely similar to the corresponding genes of man and the gorilla. These genes contain identical promoter and termination signals and have exons 1 and 2 separated by the conserved short intron 1 (122 bp) and exons 2 and 3 separated by the more rapidly evolving, larger intron 2 (893 bp and 887 bp in chimpanzee G gamma and A gamma, respectively). Each intron 2 has a stretch of simple sequence DNA (TG)n serving possibly as a "hot spot" for recombination. The two chimpanzee genes encode polypeptide chains that differ only at position 136 (glycine in G gamma and alanine in A gamma) and that are identical to the corresponding human chains, which have aspartic acid at position 73 and lysine at 104 in contrast to glycine and arginine at these respective positions of the gorilla A gamma chain. Phylogenetic analysis by the parsimony method revealed four silent (synonymous) base substitutions in evolutionary descent of the chimpanzee G gamma and A gamma codons and none in the human and gorilla codons. These Homininae (Pan, Homo, Gorilla) coding sequences evolved at one-tenth the average mammalian rate for nonsynonymous and one-fourth that for synonymous substitutions. Three sequence regions that were affected by gene conversions between chimpanzee G gamma and A gamma loci were identified: one extended 3' of the hot spot with G gamma replaced by the A gamma sequence, another extended 5' of the hot spot with A gamma replaced by G gamma, and the third conversion extended from the 5' flanking to the 5' end of intron 2, with G gamma replaced here by the A gamma sequence. A conversion similar to this third one has occurred independently in the descent of the gorilla genes. The four previously identified conversions, labeled C1-C4 (Scott et al. 1984), were substantiated with the addition of the chimpanzee genes to our analysis (C1 being shared by all three hominines and C2, C3, and C4 being found only in humans). Thus, the fetal genes from all three of these hominine species have been active in gene conversions during the descent of each species.      

“Stepping-sticks” and “seat-sticks”: New types of tools used by wild chimpanzees (Pan troglodytes) in Sierra Leone

In Tenkere, Sierra Leone, a community of wild chimpanzees (Pan troglodytes verus) spent long hours eating the fruits and flowers of the Kapok (Ceiba pentandra) tree. The branches of this species are covered in sharp thorns which make movement in their high canopies problematic for the chimpanzees. In an apparent attempt to increase their mobility and to ease the discomfort of lengthy bouts of eating in these trees, some of the Tenkere chimpanzees have been observed using stick tools as foot (“stepping-sticks”) and body (“seat-sticks”) protection against the painful thorns. This form of tool-using is culturally unique to the Tenkere chimpanzees, as at other sites where these apes have been observed eating parts of kapok trees, there are no published records of this tool technology. In three of the stepping-stick tool use incidents, the chimpanzee used the tool(s), held between their greater and lesser toes, in locomotion. This form of tool use is the first recorded case of habitually used tools that can be justifiably categorized as being “worn” by any known wild population of Pan troglodytes. Am J Primatol 41:45–52, 1997. © 1997 Wiley-Liss, Inc.