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.     

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.     

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.     

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.     

‘Impact hunters’ catalyse cooperative hunting in two wild chimpanzee communities

Even when hunting in groups is mutually beneficial, it is unclear how communal hunts are initiated. If it is costly to be the only hunter, individuals should be reluctant to hunt unless others already are. We used 70 years of data from three communities to examine how male chimpanzees ‘solve’ this apparent collective action problem. The ‘impact hunter’ hypothesis proposes that group hunts are sometimes catalysed by certain individuals that hunt more readily than others. In two communities (Kasekela and Kanyawara), we identified a total of five males that exhibited high hunt participation rates for their age, and whose presence at an encounter with red colobus monkeys increased group hunting probability. Critically, these impact hunters were observed to hunt first more often than expected by chance. We argue that by hunting first, these males dilute prey defences and create opportunities for previously reluctant participants. This by-product mutualism can explain variation in group hunting rates within and between social groups. Hunting rates declined after the death of impact hunter FG in Kasekela and after impact hunter MS stopped hunting frequently in Kanyawara. There were no impact hunters in the third, smaller community (Mitumba), where, unlike the others, hunting probability increased with the number of females present at an encounter with prey.     

Influence of Chimpanzee Predation on Associations Between Red Colobus and Red-tailed Monkeys at Ngogo,Kibale National Park,Uganda

Colobines often associate with cercopithecines at various African sites. Such polyspecific associations presumably have an antipredation function. At Ngogo, Kibale National Park, Uganda, red colobus (Procolobus rufomitratus) spend considerable time in association with red-tailed monkeys (Cercopithecus ascanius), and they are also heavily hunted by chimpanzees (Pan troglodytes). I conducted behavioral observations and playback experiments to test the hypothesis that red colobus and red-tailed monkeys obtain mutual protection and predator-related benefits by associating. Despite high chimpanzee hunting pressure on red colobus and much lower hunting pressure on red-tailed monkeys, red-tailed monkeys initiate, maintain, and terminate the associations. The results suggest that rather than providing red colobus with protection against chimpanzees, the associations occur mostly because they protect red-tailed monkeys against predation by eagles.     

Big game hunting in New Zealand: per capita effort,harvest and expenditure in 2011–2012

Recreational big game hunters make a significant contribution to conservation through kills of deer, pigs, chamois and tahr. New opportunities for managing recreational hunting through the proposed Game Animal Council underscore the need to understand the implications of potential changes in recreational hunting participation and harvests. Based on a survey of hunters' recall over a year, hunters averaged 15.63 (SEM = 0.58) big game hunts per year, spending 30.53 (SEM = 0.85) days hunting and killing 8.92 (SEM = 0.69) big game animals. Hunters commonly targeted several species on a single hunt, with highly skewed distributions for hunter effort and kills. Mean monthly expenditure on big game hunting items was $296.78 (SEM = $8.95). Results demonstrate that big game hunting is a significant activity in New Zealand, but this varies considerably among hunters with a small number responsible for the vast majority of kills. These are important considerations for future big game hunting management.     

Demographic influences on the behavior of chimpanzees

Recent research has revealed substantial diversity in the behavior of wild chimpanzees. Understanding the sources of this variation has become a central focus of investigation. While genetic, ecological, and cultural factors are often invoked to explain behavioral variation in chimpanzees, the demographic context is sometimes overlooked as a contributing factor. Observations of chimpanzees at Ngogo, Kibale National Park, Uganda, reveal that the size and structure of the unit group or community can both facilitate and constrain the manifestation of behavior. With approximately 150 individuals, the Ngogo community is much larger than others that have been studied in the wild. We have taken advantage of the unusual demographic structure of this community to document new and intriguing patterns of chimpanzee behavior with respect to hunting, territoriality, and male social relationships. Chimpanzees at Ngogo hunt often and with a considerable degree of success. In addition, male chimpanzees there frequently patrol the boundary of their territory and engage in repeated bouts of lethal intergroup aggression. By forming two distinct subgroups, male chimpanzees at Ngogo also develop social bonds above the level of dyadic pairs. While the sheer number of chimpanzees contributes to differences in hunting, patrolling, mating, and subgrouping at Ngogo, the demographic situation may also constrain behavioral interactions. At Ngogo, male chimpanzees who are closely related genetically through the maternal line do not appear to affiliate or cooperate with each other. Demographic constraints may be responsible for this finding. In this paper, I use these examples to illustrate how the demographic context affects the possible range of behavioral options open to individuals and ultimately contributes to the explanation of behavioral diversity in chimpanzees.     

First report of prey capture from human laid snare-traps by wild chimpanzees

Chimpanzees regularly hunt for meat in the wild, including both solo and group hunting; however, theft of prey from non-chimpanzee hunters, or scavenging of carcasses is extremely rare. Here we report the first observations of a novel prey capture technique by the chimpanzees in two adjacent communities in the Budongo Conservation Field Station, Uganda. In both cases blue duikers were found caught in human laid snare traps, and then retrieved by the chimpanzees. In one case the duiker was still alive when retrieved and subsequently fully consumed by the chimpanzees. In the other, the chimpanzees encountered the duiker while alive, but retrieved it soon after its death; here only a small portion was consumed. These observations are discussed in comparison to observations of chimpanzee hunting, scavenging, and their exploitation of an environment increasingly modified by human activity.     

Primate abundance along five transect lines at Ngogo, Kibale National Park, Uganda

Using the line transect methods, I studied the primate density at Ngogo, Kibale National Park, Uganda for 18 months. Comparisons with other studies show that the population of red colobus monkeys (Procolobus rufomitratus) and blue monkeys (Cercopithecus mitis) is declining, whereas the populations of black-and-white colobus (Colobus guereza), red-tailed monkeys (Cercopithecus ascanius), grey-cheeked mangabeys (Lophocebus albigena), baboons (Papio anubis), and chimpanzees (Pan troglodytes) remain constant or slightly increase. In this paper, I compare data on density from this study to data from previous and recent censuses at Ngogo and with data from other sites in the Kibale forest to examine the stability of primate populations. Furthermore, I test the hypothesis that the changes in red colobus and blue monkey density are due to changes in the forest structure and abundance of their most selected feeding trees, and show that changes in forest composition cannot account for changes in their red colobus abundance, but that hunting by chimpanzees provides a reasonable explanation.     

Interspecific interactions between wild pygmy chimpanzees (Pan paniscus) and red colobus (Colobus badius)

Interspecific interactions accompanied by physical contacts between wild pygmy chimpanzees (Pan paniscus) and red colobus (Colobus badius) were observed on three occasions at Wamba, Republic of Zaire. In all cases, the red colobus initiated the interactions by approaching the pygmy chimpanzees. Most of the pygmy chimpanzees, which were within 5 m of the red colobus, were juveniles or infants but the adult male pygmy chimpanzees never showed any interest in the red colobus. The red colobus groomed the chimpanzees in two cases, but the latter never groomed the former. No true aggressive interactions were observed between the two species. The lack of any evidence of hunting of red colobus through longitudinal studies of the pygmy chimpanzees of Wamba, together with the present observations, suggests that red colobus are probably not targets of hunting by the pygmy chimpanzees.     

Behavioral responses of male elk to hunting risk

Prey respond to predation risk with a range of behavioral tactics that can vary based on space use and hunting mode of the predator. Unlike other predators, human hunters are often more spatially and temporally restricted, which creates a period of short-duration, high-intensity predation risk for prey. Consequently, identifying the roles different hunting modes (i.e., archery and rifle), hunts for targeted and non-targeted species, and landscape features play in altering spatial and temporal responses of prey to predation risk by humans is important for effective management of harvested populations. From 2009 to 2016, we used a large-scale experiment including 50 animal-years of location data from 38 unique male elk (Cervus canadensis) to quantify changes in movement and resource selection in response to hunters during 3 separate 5-day controlled hunts for antlered males (elk archery, deer [Odocoileus spp.] rifle, and elk rifle) at the Starkey Experimental Forest and Range in northeast Oregon, USA. We evaluated competing hypotheses regarding elk responses to varying levels of prey risk posed by the different hunt types. We predicted that the strength of elk behavioral responses would increase with perceived hunter lethality (i.e., weak response to elk archery but similar response to elk and deer rifle hunts) and that prey response would be closely associated with hunter activity within the diel cycle (greater during diurnal than nocturnal hours) and across hunting seasons. Elk responses were strongest during diurnal hours when hunters were active on the landscape and were generally more pronounced during both rifle hunts than during the archery hunt (supporting our perceived lethality hypothesis). Male elk avoided open roads across all periods except during nocturnal hours of the breeding season and alternated between avoidance of areas with high canopy cover during nocturnal hours and selection during diurnal hours. In combination these patterns led to distinct distributional changes of male elk from pre-hunt to hunt periods. Patterns of male elk selection highlight the importance of managing for heterogeneous landscapes to meet a variety of habitat, harvest, hunter satisfaction, and escapement objectives.     

Primate population dynamics over 32.9 years at Ngogo, Kibale National Park, Uganda

We present census data for eight primate species spanning 32.9 years along the same transect at Ngogo, Kibale National Park, Uganda, demonstrating major changes in the composition of the primate community. Correlated with an estimated decline of ～89% in the red colobus population was an increase in encounter rates with chimpanzee parties. Our data, along with the unusually high rates of predation by chimpanzees on red colobus at Ngogo and the fact that the chimpanzee community at Ngogo is the largest ever recorded, support the conclusion that the red colobus decline was caused primarily by chimpanzee predation. This seems to be the first documented case of predation by one nonhuman primate causing the population decline in another. We evaluated disease and interspecific competition as other possible causes of the red colobus decline, but judged them to be relatively insignificant compared with predation by chimpanzees. Notable changes in encounter rates with other primate species may have resulted from forest expansion. Those for mangabeys, redtails, and black and white colobus increased significantly. Encounter rates increased for l'Hoest's monkeys too, but the increased sightings may have been an artifact of increased habituation. Sightings of blue monkey and baboon groups declined. There was no significant change in encounter rates for all species combined. The Ngogo primate community seemed to be in a nonequilibrium state, changing from one dominated by two species, a folivore (red colobus) and a frugivorous omnivore (redtails), to one dominated by three species of frugivorous omnivores (redtails, mangabeys, and chimpanzees). This study demonstrates the importance of long-term monitoring in understanding population dynamics and the role of intrinsic variables in shaping the species composition of a community.     

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.     

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.     

Market Hunting in the Taï Region,Côte d’Ivoire and Implications for Monkey Populations

We studied the effect of market hunting on primate species in the Taï National Park and adjacent forests in Côte d’Ivoire. We assessed the impact of hunting by comparing the calculated maximal reproduction rate with the current off-take rate. We assessed the average bushmeat consumption per capita/per year from weekly investigations on bushmeat available in 88 bushmeat restaurants and markets over a 12-mo period in 1999. We derived data on preferences for particular game species from interviews of 162 bushmeat consumers, 25 subsistence hunters and 3 groups of professional hunters. Hunting pressure was highest on the larger primate species such red colobus (Procolobus badius), black- and- white colobus (Colobus polykomos) and sooty mangabeys (Cercocebus atys). The amount of primate bushmeat extracted from the Taï National Park and surrounding forests was 249,229 kg in 1999. We estimated population densities using line transect surveys. By referring to current population densities we calculated the maximum production of each species using the Robinson Redford model (2001) and assuming unhunted conditions. A comparison of current harvest levels with maximum production suggests that harvest of Procolobus badius is sustainable, whereas current off-take of Colobus polykomos, Cercocebus atys, Cercopithecus diana (diana monkey) and C. campbelli (Campbell’s monkeys) exceeds sustainability by ≤3 times. We recommend that wildlife managers promote programs that encourage the production of domestic animals as a substitute for wild meat.     

The optimal foraging theory, crowding and Swedish grouse hunters

Hunters that have options to hunt in different areas should evaluate their previous hunting success when they decide where to hunt. Following optimal foraging theory for non-human predators, we investigated if hunting success and density of other hunters on the hunting area will affect the probability of return to the same area, and if such behavioural changes will result in a higher hunting success compared to hunters that change to a new area. For this purpose, we used detailed information about willow grouse (Lagopus lagopus) hunters on state-owned land in Sweden. We found support for the optimal foraging theory application on grouse hunters’ behavioural changes according to hunting success. The return rate increased with increasing hunting success, and hunters that returned to the same area also increased their success compared to hunters that changed to a new area. Only one third of the hunters returned to the same area the subsequent year. We also found a negative effect of density of hunters in an area on hunters’ return rates and their hunting success, suggesting crowding among Swedish grouse hunters.     

Impact of Commercial Hunting on Monkey Populations in the Taï region,Côte d'Ivoire1

We studied the impact of hunting on monkey species in the Taï National Park and adjacent forests in Côte d'Ivoire. The average wild meat consumption per capita per year was assessed from market surveys and interviews. We determined that the amount of primate wild meat being extracted in the Taï National Park and surrounding forests was 249 t in 1999. Hunting pressure was the highest on the larger primate species such as red colobus, Procolobus badius, black and white colobus, Colobus polykomos, and the sooty mangabey, Cercocebus torquatus atys. Estimates of population densities were based on line transect surveys. The maximum annual production of each species was calculated using the Robinson and Redford model (1991) and assuming unhunted conditions. Comparing current harvest levels with the maximum sustainable yield suggests that harvest of red colobus monkeys (Procolobus badius) is sustainable, whereas current off‐take of the black and white colobus (Colobus polykomos), the sooty mangabey (Cercocebus atys), diana monkeys (Cercopithecus diana), and Campbell's monkeys (Cercopithecus campbelli) exceeds sustainability by up to three times.     

Lionfish predators use flared fin displays to initiate cooperative hunting

Despite considerable study, mystery surrounds the use of signals that initiate cooperative hunting in animals. Using a labyrinth test chamber, we examined whether a lionfish, Dendrochirus zebra, would initiate cooperative hunts with piscine partners. We found that D. zebra uses a stereotyped flared fin display to alert conspecific and heterospecific lionfish species Pterois antennata to the presence of prey. Per capita success rate was significantly higher for cooperative hunters when compared with solitary ones, with hunt responders assisting hunt initiators in cornering the prey using their large extended pectoral fins. The initiators would most often take the first strike at the group of prey, but both hunters would then alternate striking at the remaining prey. Results suggest that the cooperative communication signal may be characteristic to the lionfish family, as interspecific hunters were equally coordinated and successful as intraspecific hunters. Our findings emphasize the complexity of collaborative foraging behaviours in lionfish; the turn-taking in strikes suggests that individuals do not solely try to maximize their own hunting success: instead they equally share the resources between themselves. Communicative group hunting has enabled Pteroine fish to function as highly efficient predators.     

Big-game,small-game,and gamebird hunting in New Zealand: Hunting effort,harvest, and expenditure in 1988

Abstract

A postal survey of 8639 licensed firearm owners in 1989 indicated that an estimated 117 200 ± 6300 New Zealanders (3.5% of the total population) did some hunting in 1988. An estimated 33 100 former hunters did not hunt in 1988 but thought it likely that they would hunt again in future. The survey provided useful estimates of 1988 national totals for hunting effort (4.4 million hunter days), gross expenditure ($NZ100 million), and harvest (6.5 million animals).

Small-game hunting dominated, involving 81% of hunters, 59% of total hunting effort, and 86% of total numerical harvest: rabbits (Oryctolagus cuniculus) and possums (Trichosurus vulpecula) each comprised 40% of the national bag. Gamebird hunting involved 48% of hunters, 19% of effort, and 11% of total harvest: ducks comprised 73% of the gamebird harvest. Although big-game hunting attracted the fewest hunters (42%) it ranked second for hunting effort (21%). Big game formed 3% of the total numerical harvest: pigs (Sus scrofa), goats (Capra hircus), and red deer (Cervus elaphus scoticus) were the most commonly taken. Big-game were estimated to comprise 49% of total harvest biomass, followed by small-game (47%), and gamebirds (4%). Most deer (nearly 60%) were taken for recreation, with helicopter-based hunting accounting for only one-third the total deer harvest.

A quarter of those people hunting in 1988 hunted on five or fewer days that year, and a relatively small group of mainly professional hunters accounted for a disproportionately large share of the overall harvest Hunters reported spending an average of $851 each on hunting in 1988. Expenditure on big-game comprised 44% of the total, small-game 23%, and gamebirds 33%. Expenditure per animal harvested or per day hunted was greater for big-game animals (other than goats) than for gamebirds, which were more expensive to hunt than small-game. Total hunting effort was inversely related to the average expenditure per animal harvested, regardless of the type of game.