Chimpanzee predation in the Mahale mountains from August 1979 to May 1982

Fifty-four episodes of predatory behavior of wild chimpanzees were recorded in Mahale, western Tanzania, from August 1979 to May 1982. The chimpanzees most frequently hunt in two seasons, during May, and from August to December. Longer-term fecal analysis indicates that predation frequency is significantly higher in the dry than in the rainy season. The seasonality of predation might be the result of the sum of various ecological factors, at least one of which is the birth season of the prey species. Most of the prey are juvenile blue duiker, bushbuck, bushpig, red colobus, and red-tailed monkeys. Sex difference is recognized in the prey selection and in the hunting method employed. Apparent local difference in the predatory behavior between Mahale and Combe chimpanzees (in Mahale,females hunt more frequently, and blue duiker is the most frequent prey) can be understood in terms of the difference either in the observation methods or in the faunal diversity and density. Other aspects of predatory behavior also are reported.     

Analysis of a bone assemblage made by chimpanzees at Gombe National Park, Tanzania

Chimpanzee hunting provides information on prey characteristics and constraints acting on a large-bodied primate lacking a hunting technology, and has important implications for modeling hunting by fossil hominids. Analysis of the remains of five red colobus monkeys captured and consumed by Gombe chimpanzees in a single hunting bout provides one of the first opportunities to investigate the characteristics of prey bones surviving chimpanzee consumption. Four of the five individuals (an older infant, two juveniles and one subadult) were preserved in the bone assemblage; a neonate was entirely consumed. Cranial and mandibular fragments had the highest survivorships, followed by the scapulae and long bones. Post-cranial axial elements had the lowest survivorships. A high percentage (80%) of the long bones and ribs surviving consumption were damaged, most commonly through crenulation and step fracturing of bone ends. One of two partially reconstructed crania preserves a canine puncture through its left parietal. Proposed characteristics of faunal assemblages formed through chimpanzee-like hunting include small modal prey size, limited taxonomic diversity, a high proportion of immature individuals and a high frequency of skull bones. These characteristics would not uniquely identify hunting by fossil primates in the geological record, necessitating a contextual approach to diagnose hunting by hominids not forming an archeological record.Hominid utilization of vertebrate tissue is first unambiguously documented at 2.5 m.y.a. Rather than representing a strict "scavenging phase" in the evolution of hominid-prey interactions, Oldowan hominid carnivory may represent the overlay of large mammal scavenging on a tradition of small mammal hunting having a low archeological visibility.     

Scavenging by chimpanzees at Ngogo and the relevance of chimpanzee scavenging to early hominin behavioral ecology

Chimpanzees regularly hunt a variety of prey species. However, they rarely scavenge, which distinguishes chimpanzee carnivory from that of some modern hunter-gatherers and, presumably, at least some Plio-Pleistocene hominins. I use observations made over an 11-year period to document all known opportunities for scavenging encountered by chimpanzees at Ngogo, Kibale National Park, Uganda, and describe all cases of scavenging. I also review data on scavenging from other chimpanzee research sites. Chimpanzees at Ngogo encountered scavenging opportunities only about once per 100 days and ate meat from scavenged carcasses only four times. Scavenging opportunities are also rare at other sites, even where leopards are present (Mahale, Ta?, Gombe), and scavenging of leopard kills is known only from Mahale. Feeding on prey that chimpanzees had hunted but then abandoned is the most common form of scavenging reported across study sites. For example, several individuals at Ngogo ate meat from a partially consumed red colobus carcass abandoned after a hunt the previous day. Such behavior probably was not common among Oldowan hominins. Ngogo data and those from other sites also show that chimpanzees sometimes eat meat from carcasses of prey that they did not see killed and that were not killed by chimpanzees, and that scavenging allows access to carcasses larger than those of any prey items. However, chimpanzees ignore relatively many opportunities to obtain meat from such carcasses. Scavenging may be rare because fresh carcasses are rare, because the risk of bacterial infections and zoonoses is high, and because chimpanzees may not recognize certain species as potential prey or certain size classes of prey species as food sources. Its minimal nutritional importance, along with the absence of technology to facilitate confrontational scavenging and rapid carcass processing, apparently distinguishes chimpanzee foraging strategies from those of at least some Oldowan hominins.     

Hunting behavior of wild chimpanzees in the Taï National Park

Hunting is often considered one of the major behaviors that shaped early hominids' evolution, along with the shift toward a drier and more open habitat. We suggest that a precise comparison of the hunting behavior of a species closely related to man might help us understand which aspects of hunting could be affected by environmental conditions. The hunting behavior of wild chimpanzees is discussed, and new observations on a population living in the tropical rain forest of the Taï National Park, Ivory Coast, are presented. Some of the forest chimpanzees' hunting performances are similar to those of savanna-woodlands populations; others are different. Forest chimpanzees have a more specialized prey image, intentionally search for more adult prey, and hunt in larger groups and with a more elaborate cooperative level than savanna-woodlands chimpanzees. In addition, forest chimpanzees tend to share meat more actively and more frequently. These findings are related to some theories on aspects of hunting behavior in early hominids and discussed in order to understand some factors influencing the hunting behavior of wild chimpanzees. Finally, the hunting behavior of primates is compared with that of social carnivores.     

Demographic influences on the hunting behavior of chimpanzees

We investigated hunting in an unusually large community of wild chimpanzees at Ngogo in the Kibale National Park, Uganda. Aspects of predation were recorded with respect to the prey, the predators, and hunting episodes. During 23 months of observation, the Ngogo chimpanzees caught 128 prey items from four primate and three ungulate species. Chimpanzees preyed selectively on immature red colobus primarily during group hunts, with adult males making the majority of kills. Party size and composition were significant predictors of the probability that chimpanzees would hunt and of their success during attempts. Chimpanzees were more likely to hunt red colobus if party size and the number of male hunters were large; party size and the number of male hunters were also significantly larger in successful compared with unsuccessful hunts. The Ngogo chimpanzees did not appear to hunt cooperatively, but reciprocal meat-sharing typically took place after kills. Hunts occurred throughout the year, though there was some seasonality as displayed by periodic hunting binges. The extremely high success rate and large number of kills made per successful hunt are the two most striking aspects of predation by the Ngogo chimpanzees. We compare currently available observations of chimpanzee hunting behavior across study sites and conclude that the large size of the Ngogo community contributes to their extraordinary hunting success. Demographic differences between groups are likely to contribute to other patterns of interpopulation variation in chimpanzee predation. Am J Phys Anthropol 109:439–454, 1999. © 1999 Wiley-Liss, Inc.     

Savanna chimpanzees, Pan troglodytes verus, hunt with tools

Although tool use is known to occur in species ranging from naked mole rats [1] to owls [2], chimpanzees are the most accomplished tool users [3-5]. The modification and use of tools during hunting, however, is still considered to be a uniquely human trait among primates. Here, we report the first account of habitual tool use during vertebrate hunting by nonhumans. At the Fongoli site in Senegal, we observed ten different chimpanzees use tools to hunt prosimian prey in 22 bouts. This includes immature chimpanzees and females, members of age-sex classes not normally characterized by extensive hunting behavior. Chimpanzees made 26 different tools, and we were able to recover and analyze 12 of these. Tool construction entailed up to five steps, including trimming the tool tip to a point. Tools were used in the manner of a spear, rather than a probe or rousing tool. This new information on chimpanzee tool use has important implications for the evolution of tool use and construction for hunting in the earliest hominids, especially given our observations that females and immature chimpanzees exhibited this behavior more frequently than adult males.     

Influence of chimpanzee predation on the red colobus population at Ngogo, Kibale National Park, Uganda

Frequent hunting of red colobus monkeys (Procolobus rufomitratus) takes place at all long-term chimpanzee (Pan troglodytes) study sites where both species are present. Red colobus are the most commonly selected prey of chimpanzees even when other monkey species are more abundant. In particular, the chimpanzee community at Ngogo, Kibale National Park, Uganda, preys heavily on red colobus monkeys: the chimpanzee hunting success rate is extremely high, and chimpanzees kill many individuals per successful hunt. Census data had suggested that the red colobus population is declining and that predation by chimpanzees may be contributing to this decline. In this paper, I address the impact of hunting on the red colobus population at Ngogo. To test the hypothesis that chimpanzee hunting is sustainable, I am using demographic data collected on red colobus monkeys over a period of 3 years, as well as fecundity and mortality data from previous studies of this species. I apply matrix models and vortex analyses using a sensitivity analysis approach to project future population development. Results show that current rates of hunting are not sustainable, but that chimpanzees are neither more “noble”, nor more “savage” than humans are, but that they also hunt to ensure maximum benefit without regard for the consequences for the prey population.     

Predation on mammals by the chimpanzee (Pan troglodytes)

With respect to prey selectivity and predation frequency, chimpanzees (Pan troglodytes) show local differences as well as diachronic variability within the same population. When data on predation from three long-term studies at Mahale, Gombe, and Tai are compared, some differences and similarities emerge; Mahale is more like Gombe than Tai in regard of prey selection but features of hunting at Tai with respect to predation frequency are not conspicuous. The most responsible factor for diversity in prey selectivity is a distinct “prey image” maintained by chimpanzees of different populations, although it is necessary to clarify in future studies why and how such tradition develops. Relative body size of chimpanzees to prey species and/or the degree of cooperation among members of a hunting party may explain the variability in prey size selected at each site, the latter influencing the frequency of successful hunts at the same time. Although various degrees of habituation and different sampling methods including artificial feeding might have obscured the real differences, recent data from the three populations do not seem to be biased greatly by such factors. Nevertheless, it is still difficult to make strict comparisons due to the lack of sufficient standardized data across the three populations on the frequency of hunting and predation. It is suggested that the size or demographic trend of a chimpanzee unit-group, especially the number of adult males included, necessarily influences its hunting frequency as well as its prey profile. It is also suggested that factors which bring these males together into a party (e.g. fruit abundance, swollen females, conflict between unit-groups etc.) strongly affect theactual hunting and kill rates. Other possible factors responsible for the local differences are forest structure (e.g. tree height), skilful “hero” chimpanzees, and competition with sympatric carnivorous animals. A total of at least 32 species have been recorded as prey mammals of chimpanzees from 12 study sites and the most common prey mammals are primates (18 species), of which 13 species are forest monkeys. Forest monkeys, colobine species in particular, are often the most common victims of the predation by chimpanzees at each site. We may point out a tendency toward selective hunting for the forest monkeys in terms of the selectivity of prey fauna among all three subspecies of chimpanzees, including populations living in drier environment. The mode of chimpanzee hunting seems to correspond to the highest available biomass of gregarious, arboreal monkeys in the forest, colobine species in particular. In contrast, bonobos (P. paniscus) are less carnivorous than chimpanzees, only rarely preying on a few species of small mammals. The sharp contrast of the two allied species in their predatory tendencies appears to have something to do with the differences in the structure of primary production between their habitats.     

Cooperative hunting roles among taï chimpanzees

All known chimpanzee populations have been observed to hunt small mammals for meat. Detailed observations have shown, however, that hunting strategies differ considerably between populations, with some merely collecting prey that happens to pass by while others hunt in coordinated groups to chase fast-moving prey. Of all known populations, Taï chimpanzees exhibit the highest level of cooperation when hunting. Some of the group hunting roles require elaborate coordination with other hunters as well as precise anticipation of the movements of the prey. The meat-sharing rules observed in this community guarantee the largest share of the meat to hunters who perform the most important roles leading to a capture. The learning time of such hunting roles is sometimes especially long. Taï chimpanzee males begin hunting monkeys at about age 10. The hunters’ progress in learning the more sophisticated hunting roles is clearly correlated with age; only after 20 years of practice are they able to perform them reliably. This lengthy learning period has also been shown in some hunter-gatherer societies and confirms the special challenge that hunting represents.     

Cockscomb Revisited: Jaguar Diet in the Cockscomb Basin Wildlife Sanctuary,Belize1

The Cockscomb Basin Wildlife Sanctuary, Belize was established in 1986 following Rabinowitz and Nottingham's 1986 study on jaguar diet and distribution. In 2002, we reexamined jaguar diet following two decades of mitigated human hunting of jaguar prey. We observed high overlap between historical and current jaguar diet constructs. Nine‐banded armadillo remained the most frequently consumed prey item. However, collared peccary consumption increased by nearly 20 percent while dietary breadth decreased by 60 percent presumably reflecting increased prey availability following reduced human hunting pressure.     

Use of tool sets by chimpanzees for multiple purposes in Moukalaba-Doudou National Park,Gabon

We report our recent findings on the use of tool sets by chimpanzees in Moukalaba-Doudou National Park, Gabon. Direct observations and evidences left by chimpanzees showed that chimpanzees used sticks as pounders, enlargers, and collectors to extract honey from beehives of stingless bees (Meliponula sp.), which may correspond to those previously found in the same site for fishing termites and to those found in Loango National Park, Gabon. However, we observed chimpanzees using a similar set of tools for hunting a medium-sized mammal (possibly mongoose) that hid inside a log. This is the first report of hunting with tools by a chimpanzee population in Central Africa. Chimpanzees may recognize the multiple functions and applicability of tools (extracting honey and driving prey), although it is still a preliminary speculation. Our findings may provide us a new insight on the chimpanzee’s flexibility of tool use and cognitive abilities of complex food gathering.     

THE EFFECTS OF TSETSE CONTROL OPERATIONS ON COMMON DUIKER IN EASTERN ZAMBIA

The effects of intensive Tsetse Control hunting operations on a duiker population (Sylvicapra grimmia) were investigated in a 200 sq. mile area in Eastern Zambia. Two years of hunting were insufficient to reduce this population so significantly that a marked shift of its age composition towards the juvenile age classes resulted. There were, however, indications of beginning accelerated population growth through increased breeding and inclusion of more juveniles in the reproduction process, as a first response to the hunting pressure. Although general availability of duiker did not diminish, they became increasingly difficult to shoot because of behavioural adaptation and changing periods of feeding activity. Neither hunting nor various other human disturbances provoked emigration from the area or a change of the seasonal pattern of localised movement. The studied hunting operation failed to remove more than the annual increment to the duiker population and in respect of this species was thus ineffective as a means of Tsetse Control. Implications of the results of this study for the management of duiker for sustained meat production are discussed.     

The Influence of Hunting on Antipredator Behavior in Central African Monkeys and Duikers

Many animals can adjust their behavioral strategies to reduce predation risk. We investigated whether rain forest monkeys and duikers alter their antipredatory behavior in response to hunting by humans in southwestern Gabon. We compared monkey and duiker responses to human observers in an area where hunting is prohibited, to those in a nearby area where hunting pressure is moderate but spatially variable. The results of our study indicate that monkeys become more secretive when hunted, commencing alarm calls only when at a certain distance (typically > 50 m) from humans. We found no difference in monkey group size between hunted and no-hunting areas. In no-hunting areas, duikers often freeze in response to approaching observers, but in hunted areas they abandon this strategy and rapidly flee from humans. Duikers also whistle more often in areas where they are hunted frequently. Our findings have at least two important implications. First, behavioral observations of monkeys and duikers may be useful in gauging local hunting intensity in African rain forests. Second, duiker densities are likely to be overestimated in hunted areas, where they more readily flee and whistle, and underestimated in no-hunting areas, where they rely on freezing behavior to avoid detection. Because behavioral adaptations to hunting vary both among species and localities, these differences should be considered when attempting to derive population-density estimates for forest wildlife.     

Feeding behaviour of the bats Nycteris grandis and Nycteris thebaica (Nycteridae) in captivity

The feeding and hunting behaviour of Nycteris grandis and N. thebaica was observed in captivity at the Sengwa Wildlife Research Area in Zimbabwe in January and February 1982. Both species preferentially selected katydids and beetles over moths, and relied heavily on acoustic stimuli emanating from prey to detect targets. Nycteris grandis readily consumed frogs and bats and appeared not to use the calls of male frogs or the echolocation calls of other bats to locate prey. Both species produced echolocation calls during attacks on prey, increasing the rates of pulse repetition as they closed with targets and suggesting the use of echolocation in hunting. The echolocation calls of N. grandis are described along with general observations of the behaviour of both species.     

Capturing and toying with hyraxes (Dendrohyrax dorsalis) by wild chimpanzees (Pan troglodytes) at Bossou, Guinea

Chimpanzees (Pan troglodytes verus) were observed capturing and toying with western tree hyraxes (Dendrohyrax dorsalis, Order Hyracoidea) at Bossou, Guinea. An adolescent female carried one hyrax for 15 hr, slept with it in her nest, and groomed it. The captive was not consumed. Nearby adults ignored the hyrax. In another case, two adolescent males timidly inspected a small hyrax. These observations indicate that the chimpanzees at Bossou do not regard the hyrax as a prey animal, supporting the idea that lack of opportunity does not seem to be the only reason that chimpanzees do not consume an individual of a potential prey species.     

Night and day: evaluating transect methodologies to monitor duikers in the Dzanga‐Sangha Protected Areas,Central African Republic

Across the Congo Basin, human hunting pressure poses the biggest threat to small‐ and medium‐bodied forest ungulates (genus Philantomba and Cephalophus, commonly known as duikers). The exploitation of these species has cascading effects on larger ecosystem processes, as well as on human subsistence practices. This study compares encounter rates and estimated population densities of duiker species, specifically Philantomba monticola (blue duiker) in the Dzanga‐Sangha Protected Areas (APDS), Central African Republic (CAR). Data were collected using direct observations of individual animals during diurnal (135 km) and nocturnal (150 km) transects in the APDS, with abundance estimates produced using DISTANCE software. Transect data demonstrate that within hunted forests similar to APDS, nocturnal rather than diurnal transects yield more individual observations of ungulates. Despite hunting pressure in the region, estimates presented for APDS suggest some of the highest density estimates reported for blue duikers in western and central Africa (58.6 blue duikers per km²). This study directly contributes current regional data on the status of duiker populations at APDS and in the larger Sangha Trinational Landscape (TNS, UNESCO). More broadly, we highlight the potential importance of nocturnal transect data to the development of adaptive management regimes in hunted forests.     

New records on prey capture and meat eating by bonobos at Lui Kotale, Salonga National Park, Democratic Republic of Congo

Compared to data from chimpanzees, observations on prey capture and meat eating by bonobos (Pan paniscus) are still rare, fragmentary and anecdotal. Here we present new and unpublished information from wild bonobos at Lui Kotale, Salonga National Park, Democratic Republic of Congo. Our observations confirm that solitary and terrestrial ungulates are the major prey. However, bonobos at Lui Kotale also consumed other mammalian prey, including other primates. Evidence from direct observations is complemented with information obtained by macroscopic analyses of fresh faeces. Results suggest that bonobos consume meat with frequencies similar to some chimpanzee populations. The data emphasize differences between the two Pan species in terms of prey species selection and prey capture.     

许家窑遗址马科动物的死亡年龄

普氏野马(Equus przewalskii)和野驴(Equus hemionus)是许家窑遗址动物群中的优势属种。本文基于对这两种动物牙齿材料的测量与分析,确定了遗址中马科动物的死亡年龄,并对上、下文化层的死亡年龄分布进行了研究,以期探知古人类获取肉食资源的方式与特点。通过与马科动物在自然生存状态下以及死于不同原因(如疾病或营养衰竭、食肉动物猎杀、现代人类狩猎等)的年龄结构对比,结果表明:古人类在许家窑文化早期(下文化层)可能通过捡拾自然死亡的动物尸体、与食肉类动物抢夺猎物、主动狩猎等多种方式获取马科动物,而在许家窑文化晚期(上文化层)可能以主动狩猎作为获取马科动物的主要方式。此外,古人类在遗址的早期就可能已经具有捕获整个马科动物居群中任意年龄个体的能力,并能做出最优化判断,有选择地去捕猎脂肪和肉量较高的壮年动物群体。     

Patterns of predation by chimpanzees on red colobus monkeys in gombe national park, 1982–1991

Predatory patterns in wild chimpanzees are important evidence in the continuing debate about the role of hunting in the behavior of early hominids. Data are presented on the predator–prey ecology of red colobus monkeys (Colobus badius tephrosceles) and chimpanzees (Pan troglodytes schweinfurthii) in Gombe National Park, Tanzania, from 1982 through 1991. During this period chimpanzees were observed to kill 429 mammalian prey items, 350 of which were red colobus. Hunts were undertaken by chimpanzees in 71.5% of encounters with red colobus, and in 52.2% of all hunts at least one colobus was caught. Hunting occurred in all months, but its frequency peaked in the late dry season months of August and September, and was lowest in the rainy months of April and May. There was greater seasonality of hunting from 1982 to 1991 than previously reported for Gombe. Hunting success varied between 40% in the rainy season and 65% in the dry season. Sixty multiple kills of colobus were reported in which from two to seven colobus were killed. Approximately 75% of all colobus caught were immatures; juveniles were the most preyed upon age class. Adult and adolescent male chimpanzees made 89.3% of all kills; the 10.7% of kills made by adult females was an increase over the 4% figure for female kills reported in the preceding decade. Hunting showed a strong “binge” tendency, with the explanation for binges likely related to social rather than ecological factors. These results are discussed in light of earlier hunting data for Gombe chimpanzees, and compared with data from other chimpanzee field studies. © 1994 Wiley-Liss, Inc.