Hunting, law enforcement, and African primate conservation

Abstract: Primates are regularly hunted for bushmeat in tropical forests, and systematic ecological monitoring can help determine the effect hunting has on these and other hunted species. Monitoring can also be used to inform law enforcement and managers of where hunting is concentrated. We evaluated the effects of law enforcement informed by monitoring data on density and spatial distribution of 8 monkey species in Taï National Park, Côte d’Ivoire. We conducted intensive surveys of monkeys and looked for signs of human activity throughout the park. We also gathered information on the activities of law‐enforcement personnel related to hunting and evaluated the relative effects of hunting, forest cover and proximity to rivers, and conservation effort on primate distribution and density. The effects of hunting on monkeys varied among species. Red colobus monkeys (Procolobus badius) were most affected and Campbell's monkeys (Cercopithecus campbelli) were least affected by hunting. Density of monkeys irrespective of species was up to 100 times higher near a research station and tourism site in the southwestern section of the park, where there is little hunting, than in the southeastern part of the park. The results of our monitoring guided law‐enforcement patrols toward zones with the most hunting activity. Such systematic coordination of ecological monitoring and law enforcement may be applicable at other sites.     

Risk-prone hunting by chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>) increases during periods of high diet quality

Most studies suggest that during times of nutritional stress, an animal faced with two foraging choices should follow a risk-prone strategy, choosing the option with highest payoff variance. This “scarcity/risk” hypothesis was developed to account for the foraging patterns of small animals with high metabolic rates susceptible to the threat of starvation. In this paper, we propose that animals should also be risk-prone when their diet quality is particularly high, far exceeding that which is needed to survive. Under these circumstances, the costs of experiencing a low or negative payoff can easily be recouped. We suggest that large-bodied omnivores are most likely to adopt this “abundance/risk” strategy. We investigate this question among wild chimpanzees (Pan troglodytes) that choose between a risk-averse strategy of feeding on plant material and a risk-prone strategy of hunting red colobus monkeys. Using 14 years of data on the Kanyawara chimpanzees of Kibale National Park, Uganda, we find strong evidence that chimpanzees follow the “abundance/risk” strategy. Both hunting rate (hunts/100 observation hours) and the probability of hunting upon encountering red colobus monkeys were positively correlated with seasonal consumption of ripe drupe fruits, a class of preferred food associated with elevated reproductive performance by females. Critically, these results remained statistically significant after controlling for the potentially confounding effects of male chimpanzee party size and the presence of sexually receptive females. These findings suggest that the relationship between risk-sensitive foraging and diet quality depends upon the daily probability of starvation, the number of alternative foraging strategies, and the degree to which diet quality satisfies an animal’s nutritional requirements.     

Female gregariousness in Western Chimpanzees (Pan troglodytes verus) is influenced by resource aggregation and the number of females in estrus

Among social animals, group size is constrained by competition over resources. Because female reproductive success is limited by access to food resources, and that of males by access to fertile females, chimpanzee females are proposed to be less social than males and to maintain weak intrasexual relations. Findings from Taï National Park, Côte d’Ivoire, challenged this view, as chimpanzee females were described as generally gregarious, and close intrasexual bonds were common. Here, in a new analysis that focuses on the South Group of chimpanzees in Taï forest, we reevaluate the proposed differences in female association patterns between the Taï and East African populations. We find that mean party size and dyadic association index between females has decreased in Taï, although the level of dyadic associations remains high compared with East African chimpanzees. We attribute the decrease in female gregariousness to the decline in community size over the last 10 years. In addition, we use a multivariate approach to analyze social and ecological factors influencing party size in females. We show that female gregariousness increased when the fruit resources were more clumped and with increased number of females in estrus present. Party size of mothers with sons, however, was smaller with increasing number of sexually receptive females. The results of our model and the reviewed findings of other studies support the socioecological model because food distribution affects female gregariousness, but social and demographic aspects are equally influencing female grouping tendencies.     

Predation affects alarm call usage in female Diana monkeys (Cercopithecus diana diana)

Diana monkeys produce acoustically distinct calls to a number of external events, including different types of predators. In a previous study, we found population-wide differences in male alarm call production in Taï Forest, Ivory Coast, and on Tiwai Island, Sierra Leone, mostly likely originating from differences in predator experience. In Taï Forest, leopards (Panthera pardus) are common but on Tiwai Island they have been absent for decades, while the predation pressure from crowned eagles (Stephanoaetus coronatus) has been similar. To further evaluate the impact of predator experience, we here analyse the vocal behaviour of female Diana monkeys in both habitats. Female Diana monkeys produce predator-specific alarm calls, alert calls and contact calls in response to predators, suggesting that their calls serve in a broader range of functions compared to males. Results showed that females produced the same call types at both sites, despite the differences in predator fauna. Regarding call usage, leopard alarm calls were extremely rare on Tiwai Island, but not in Taï Forest, whereas we found no differences in eagle alarm call production. When comparing response latencies, Tiwai females were slower to respond to both predators compared to Taï females. Finally, we found no habitat-specific acoustic differences in the alert and predator-specific alarm calls, but significant differences in frequency-based parameters of contact calls. Overall, our results suggest that ontogenetic experience can affect primate vocal behaviour in both usage and acoustic structure but that the way in which particular call types are affected may be closely linked to function.     

Reciprocity and trades in wild West African chimpanzees

Why do animals help other individuals and provide benefits to the recipient, sometimes at personal cost? In this study, we aim to determine if some of the helpful behaviors observed in a group of wild chimpanzees (Taï chimpanzee group, Côte d'Ivoire, West Africa) are exchanged among individuals resulting in a net benefit for both participants. We adopted an inclusive view of exchanges by considering that all commodities (i.e., social behaviors as grooming, sex, support, as well as resources, such as meat) can be exchanged. This would result in “market” type social interactions in which debts of one commodity can be met by giving the same or other commodities. We investigated whether both in-kind exchanges and trades of commodities occurred. The Taï chimpanzees reciprocated the amount of grooming they received and were more likely to give support in agonistic conflicts to those who also supported them. In addition, they traded support for meat and meat for sex. Both male and female chimpanzees exchanged many different commodities, which seemed to result in long-term balanced relationships. Our results suggest that wild chimpanzees rely on other group members to obtain many of the important commodities they need.     

Interdemic variation in mixed-species association patterns: common diurnal primates of Kibale National Park, Uganda

We used interdemic variation in the tendency to form mixed-species groups to examine the costs and benefits of association among the primates of Kibale National Park, Uganda. A year-long survey of six sites revealed that the amount of time that the five common diurnal primates [red colobus (Procolobus tephrosceles), black-and-white colobus (Colobus guereza), redtail monkeys (Cercopithecus ascanius), blue monkeys (Cercopithecus mitis), and grey-cheeked mangabeys (Lophocebus albigena)] spent in mixed-species groups varied dramatically among sites. In many cases, the proportion of time that species associated was positively related to their densities. By using detailed behavioral observations of redtail monkeys and red colobus made over 4 years (2660 h) at four sites, we were able to reject the null hypothesis that associations occur by chance for only one of four sites. However, a correlative approach exploring the costs and benefits of association suggests that ecological variables do influence association patterns. We found that redtail monkeys and red colobus overlapped in diet (19.2% of their foraging effort) and traveled further when in mixed-species groups than when alone. Having demonstrated this, we examined the applicability of the ecological constraints model for predicting the proportion of the time spent in mixed-species groups based on food availability. For this analysis we concentrated on red colobus from the site with 35 months of observation and demonstrated that their tendency to be in mixed- species groups was related to food availability. We used two methods to examine if mixed-species associations function to decrease predation risk. First, chimpanzees are known to prey heavily on red colobus, but rarely kill other primates. The time red colobus spent in mixed-species groups was correlated to chimpanzee density, but it was not for the other monkey species, suggesting that mixed-species groups serve to decrease predation risk. Second, when red colobus groups contain more infants and are presumably at the greatest risk of predation, they form mixed-species groups most often. These results demonstrate that the costs and benefits of mixed-species associations vary dramatically over small spatial and temporal scales. If such variation is generally the case, then studies conducted at different locations or different times could easily highlight the importance of difference selective agents in favoring mixed-species associations. Received: 10 February 1999 / Received in revised form: 16 September 1999 / Accepted: 2 October 1999     

The Contribution of Long-Term Research at Gombe National Park to Chimpanzee Conservation

Abstract:  Long-term research projects can provide important conservation benefits, not only through research specifically focused on conservation problems, but also from various incidental benefits, such as increased intensity of monitoring and building support for the protection of an area. At Gombe National Park, Tanzania, long-term research has provided at least four distinct benefits to wildlife conservation. (1) Jane Goodall's groundbreaking discoveries of chimpanzee (Pan troglodytes) tool use, hunting, and complex social relationships in what was then a game reserve drew attention to the area and created support for upgrading Gombe to national park status in 1968. (2) The highly publicized findings have earned Gombe and Tanzania the attention of a worldwide public that includes tourists and donors that provide financial support for Gombe, other parks in Tanzania, and chimpanzee conservation in general. (3) Crucial information on social structure and habitat use has been gathered that is essential for effective conservation of chimpanzees at Gombe and elsewhere. (4) A clear picture of Gombe's chimpanzee population over the past 40 years has been determined, and this has helped identify the greatest threats to the viability of this population, namely disease and habitat loss outside the park. These threats are severe and because of the small size of the population it is extremely vulnerable. Research at Gombe has led to the establishment of conservation education and development projects around Gombe, which are needed to build local support for the park and its chimpanzees, but saving these famous chimpanzees will take a larger integrated effort on the part of park managers, researchers, and the local community with financial help from international donors.     

The efficacy of female choice in chimpanzees of the Taï Forest, Côte d’Ivoire

Male–male competition has historically been considered the major force driving sexual selection. However, female choice and inter-sexual conflict are increasingly recognized as important influences affecting differential mating and reproductive success. Many females exhibit preferences for particular males; however, male strategies may conflict with females’ ability to obtain their mate preferences. To influence paternity, females must affect both (1) whether or not sexual interactions occur, particularly during the periovulatory period (POP) and (2) the outcome of sexual interactions. This study focuses on the effectiveness of female choice in wild chimpanzees (Pan troglodytes verus). Over 2,600 h of data were collected on two habituated chimpanzee communities in the Taï National Park, Côte d’Ivoire. Female mate preferences were measured by quantifying proceptive and resistance behavior toward males in both the periovulatory period and non-POP phases of estrus. The efficacy of female preference was measured both (1) by measuring success rates of female proceptivity and resistance behaviors and (2) by determining how well measures of female mate preference (proceptivity and resistance rates) predict male mating success. Though male chimpanzees are clearly dominant to females, the results indicate that females could effectively resist male solicitations and, in most cases, unwanted copulations were averted. Both female proceptivity and resistance rates correlate (positively and inversely, respectively) with male mating success in POP. Outside POP, female proceptivity rates corresponded with male mating success, but resistance rates did not. Males (irrespective of rank) that were preferred by females obtained higher mating success compared to other males during the POP, suggesting that females were effective in their mate choice and that, despite clear male dominance, female choice influences paternity in wild chimpanzees.     

The meat-scrap hypothesis: small quantities of meat may promote cooperative hunting in wild chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

A common explanation for hunting in groups is that doing so yields a greater per capita caloric benefit than hunting solitarily. This is logical for social carnivores, which rely exclusively on meat for energy, but arguably not for omnivores, which obtain calories from either plant or animal matter. The common chimpanzee, Pan troglodytes, is one of the few true omnivores that regularly hunts in groups. Studies to date have yielded conflicting data regarding the payoffs of group hunting in chimpanzees. Here, we interpret chimpanzee hunting patterns using a new approach. In contrast to the classical assumption that hunting with others maximizes per capita caloric intake, we propose that group hunting is favored because it maximizes an individual’s likelihood of obtaining important micronutrients that may be found in small quantities of meat. We describe a mathematical model demonstrating that group hunting may evolve when individuals can obtain micronutrients more frequently by hunting in groups than by hunting solitarily, provided that group size is below a certain threshold. Twenty five years of data from Gombe National Park, Tanzania are consistent with this prediction. We propose that our ‘meat-scrap’ hypothesis is a unifying approach that may explain group hunting by chimpanzees and other social omnivores.     

An agent-based model of red colobus resources and disease dynamics implicates key resource sites as hot spots of disease transmission

The effect of anthropogenic landscape change on disease in wildlife populations represents a growing conservation and public health concern. Red colobus monkeys (Procolobus rufomitratus), an endangered primate species, are particularly susceptible to habitat alteration and have been the focus of a great deal of disease and ecological research as a result. To infer how landscape changes can affect host and parasite dynamics, a spatially explicit agent-based model is created to simulate movement and foraging of this primate, based on a resource landscape estimated from extensive plot-derived tree population data from Kibale National Park, Uganda. Changes to this resource landscape are used to simulate effects of anthropogenic forest change. With each change in the landscape, disease outcomes within the simulated red colobus population are monitored using a hypothetical microparasite with a directly transmitted life cycle. The model predicts an optimal distribution of resources which facilitates the spread of an infectious agent through the simulated population. The density of resource rich sites and the overall heterogeneity of the landscape are important factors contributing to this spread. The characteristics of this optimal distribution are similar to those of logged sections of forest adjacent to our study area.     

Prey selectivity by crowned hawk-eagles on monkeys in the Kibale Forest,Uganda

Summary Results are presented from a 3.25-year study of a nesting pair of crowned hawk-eagles and an 18-year study of a primate community in the Kibale Forest, Uganda. The proportional composition of the living population of prey species was compared with that of eagle prey and animals dying from other causes. Monkeys were the predominant pry (83.7%). They also dominated the medium-to large-sized mammalian carcasses dying from other causes (88.9%). The eagles selected prey according to species, age, and sex. Selectivity by age and sex differed between prey species. Among red colobus monkeys, the eagles selected young juveniles and infants, but in four other monkey species they selected adult males. Eagle prey selectivity by species generally supports the hypothesis that polyspecific associations among the monkeys are effective deterrents against predation. The prey/predator ratio for the Kibale eagles was much higher, but the annual offtake of prey by the eagles was much lower than that of tropical felids. Mortality due to causes other than eagles was greater than expected in red colobus and less in redtails, but not significantly different from expected or equivocal in the other three monkey species. Other cause of mortality affected adult male and infant red colobus more than expected. Among the other four monkey species, significantly more adult males and fewer adult females died from these other causes than expected. Eagle predation had a major impact on the populations of adult males of both black and white colobus and blue monkeys and on both adult male and female mangabeys. The selectivity appeared to contribute significantly to the differential adult sex ratio in four monkey species, but not in red colobus. In contrast, mortality incurred during fighting among adult males probably accounted for the differential adult sex ratio in red colobus. Offprint requests to: T.T. Struhsaker at his current address     

Different escape tactics of two vole species affect the success of the hunting predator, the least weasel

In the ongoing evolutionary arms race between predators and their prey, successful escape from the predator leads to the evolution of improved escape tactics in prey, but also predators become more effective in following and attacking the prey. Antipredatory behavior of prey is considered to be the strongest towards their most dangerous predators. However, prey species can differ both in vulnerability and efficiency of escape to a shared predator. We studied escape reactions of two vole species, the bank vole (Myodes glareolus) and the field vole (Microtus agrestis), under a simulated predation risk of the least weasel (Mustela nivalis nivalis). We conducted a laboratory experiment where a vole was given a possibility to escape from a weasel by fleeing to a horizontal tunnel or climbing the tree. Subsequently to the vole escape decision, we released a weasel to the same tunnel system to test how the weasel succeeded in following the vole. Weasel presence changed the behavior of voles as especially bank voles escaped by climbing. Instead, the majority of field voles fled into the ground-layer tunnel. The different escape tactics of the voles affected the success of the weasel, because climbing voles were less often successfully followed. We suggest that the difference in escape tactics has evolved as an adaptation to different habitats; meadow-exploiting field voles using ground-level escape while bank voles living in three-dimensional forest habitat frequently use arboreal escape tactics. This is likely to lead to different habitat-dependent vulnerabilities to predation in Microtus and Myodes vole species.     

Primate Conservation along the Tana River, Kenya: An Examination of the Forest Habitat

The Tana River National Primate Reserve, Kenya, was established in 1976 to preserve the endemic and endangered Tana River red colobus ( Colobus badius rufomitratus ) and crested mangabey ( Cercocebus galeritus galeritus ). Between 1975 and 1985 their populations declined by 80% and 45%, respectively. A study addressing primate-to-habitat relationships was conducted in 12 forest areas to determine what attributes appear most important to the in situ preservation of both endangered primates. Correlations among the abundances of primate groups, their group sizes, and the structural, resource, disturbance, and spatial attributes of the forest were used to distinguish the quality of forest patches. Intraforest habitat quality was examined, using canonical variate analysis to discriminate primate ranging patterns based on canopy and subcanopy tree composition. The colobus and mangabey show positive relationships to interior-forest habitat and appear susceptible to forest disturbances that reduce forest area or that increase forest edge and intraforest disturbance. Forest loss, fragmentation, and consequent reduction in the area-to-perimeter ratio of the remaining forests measured from 1960 to 1975 provide a partial explanation for the decline in primate populations. The results suggest that a combination of primary food items and seasonal food resources in large, high-stature, closed-canopy forests is the best predictor of high-quality habitat for these monkeys. Stewardship should be directed at the preservation of these areas or the restoration of forests toward this habitat model     

Towards an ecological solution to the folivore paradox: patch depletion as an indicator of within-group scramble competition in red colobus monkeys (Piliocolobus tephrosceles)

A number of socioecological models assume that within-group food competition is either weak or absent among folivorous primates. This assumption is made because their food resources are presumed to be superabundant and evenly dispersed. However, recent evidence increasingly suggests that folivore group size is food-limited, that the primates prefer patchily distributed high-quality foods, and display some of the expected responses to within-group scramble competition. To investigate this apparent contradiction between theoretical models and recent empirical data, we examined the foraging behaviour of red colobus monkeys (Piliocolobus tephrosceles) in Kibale National Park, Uganda. We found that red colobus monkeys foraged in a manner that suggests they deplete patches of preferred foods: intake rate slowed significantly during patch occupancy while movement rate, an index of foraging effort, increased. Furthermore, patch occupancy was related to the size of the feeding group and the size of the patch. These results suggest that within-group scramble competition occurs, may limit folivore group size, and should be considered in models of folivore behavioural ecology.     

Social behavior of free-ranging juvenile sooty mangabeys (Cercocebus torquatus atys)

Sooty mangabeys are terrestrial monkeys exhibiting female philopatry and male dispersal. Studies in captivity as well as in the wild have found that adult females form linear dominance hierarchies. However, while captive studies found no evidence for a matrilineal social system, a previous study in Taï National Park, Ivory Coast, suggested that relatedness could influence both dominance rank and affiliation pattern among adult females. Here I test whether the dominance rank, coalitionary behavior, and affiliative behavior of juveniles in a group of free-ranging mangabeys in the Taï National Park are in accordance with a matrilineal, individual, or age-related dominance system. I found that juvenile females' dominance ranks remained stable over time and were highly correlated with the dominance ranks of their mothers, whereas juvenile males' dominance ranks were initially correlated with the ranks of their mothers, but showed greater instability with increasing age. Moreover, coalitions occurred mainly between juveniles and animals that were close in rank, including their mothers and siblings. Finally, juvenile females associated and groomed preferentially with close-ranking juvenile and adult females. Juvenile males showed similar preferences in affiliation with adult females, but when associating with juvenile males, they preferred peers. The observed social behavior of free-ranging juvenile sooty mangabeys resembled the social behavior described for juveniles of many matrilineal primate species.     

Extinction of a West African Red Colobus Monkey

Abstract: We provide the first documented case of the extinction in the twentieth century of a widely recognized primate taxon. During surveys in Ghana and Côte d'Ivoire in 1993–1999, we were unable to find any surviving populations of Miss Waldron's red colobus monkey (   Procolobus badius waldroni ), a primate taxon endemic to the forests of this part of West Africa. We conclude that this monkey, which at least one authority considers worthy of species status, is probably extinct. Hunting by humans appears to be the ultimate cause of the extinction. Until our surveys began, little attention had been paid to the plight of this red colobus monkey, despite its listing as endangered by the World Conservation Union. The extinction of other large animals in the Upper Guinea rainforest region is likely to follow soon unless more attention is paid to the full range of endangered forms and more resources are devoted to their rigorous protection.     

The alarm call system of wild black-fronted titi monkeys, <Emphasis Type="Italic">Callicebus nigrifrons</Emphasis>

Upon encountering predators, many animals produce specific vocalisations that alert others and sometimes dissuade the predators from hunting. Callicebus monkeys are known for their large vocal repertoire, but little is known about the function and meaning of most call types. We recorded a large number of natural predator responses from five different groups of black-fronted titi monkeys in their Atlantic forest habitat in South Eastern Brazil. When detecting predatory threats, adult group members responded with call sequences that initially consisted of two brief, high-pitched calls with distinct frequency contours. Call A was mainly given to raptors but also to predatory capuchin monkeys and other threats within the canopy, while call B was given to predatory or non-predatory disturbances on the ground. In later parts of the sequences, we also recorded a high-pitched unmodulated call C and various low-pitched loud calls. Results therefore suggest that calls A and B provide listeners with rapid and reliable information about the general classes of danger experienced by the caller, while obtaining more specific information through other call types and combinations and behavioural responses. We discuss these findings in relation to current evolutionary theory of primate communication.     

Modeling the impacts of hunting on the population dynamics of red howler monkeys (Alouatta seniculus)

Overexploitation of wildlife populations occurs across the humid tropics and is a significant threat to the long-term survival of large-bodied primates. To investigate the impacts of hunting on primates and ways to mitigate them, we developed a spatially explicit, individual-based model for a landscape that included hunted and un-hunted areas. We used the large-bodied neotropical red howler monkey (Alouatta seniculus) as our case study species because its life history characteristics make it vulnerable to hunting. We modeled the influence of different rates of harvest and proportions of landscape dedicated to un-hunted reserves on population persistence, population size, social dynamics, and hunting yields of red howler monkeys. In most scenarios, the un-hunted populations maintained a constant density regardless of hunting pressure elsewhere, and allowed the overall population to persist. Therefore, the overall population was quite resilient to extinction; only in scenarios without any un-hunted areas did the population go extinct. However, the total and hunted populations did experience large declines over 100 years under moderate and high hunting pressure. In addition, when reserve area decreased, population losses and losses per unit area increased disproportionately. Furthermore, hunting disrupted the social structure of troops. The number of male turnovers and infanticides increased in hunted populations, while birth rates decreased and exacerbated population losses due to hunting. Finally, our results indicated that when more than 55% of the landscape was harvested at high (30%) rates, hunting yields, as measured by kilograms of biomass, were less than those obtained from moderate harvest rates. Additionally, hunting yields, expressed as the number of individuals hunted/year/km², increased in proximity to un-hunted areas, and suggested that dispersal from un-hunted areas may have contributed to hunting sustainability. These results indicate that un-hunted areas serve to enhance hunting yields, population size, and population persistence in hunted landscapes. Therefore, spatial regulation of hunting via a reserve system may be an effective management strategy for sustainable hunting, and we recommend it because it may also be more feasible to implement than harvest quotas or restrictions on season length.     

Dawn chorusing in guereza colobus monkeys

Dawn chorusing by guereza black-and-white colobus monkeys is one of the most impressive spectacles of African rainforests. This vocal behaviour is highly contagious, travelling from one neighbouring group to the next, until a wide forest area is covered by calling monkeys. Chorusing usually occurs up to 2 h before dawn, sometimes more than once, unless the preceding night was cold and wet. We conducted a series of playback experiments, which showed that guerezas’ chorusing was difficult to elicit experimentally, suggesting that callers took several variables into account before responding to other monkeys’ predawn calls. Acoustic analyses showed that morning calls were individually distinct, but we found no evidence that callers took individual identity into account in their decision to participate in chorusing. The only way to reliably elicit chorusing in our study area was to broadcast recordings of morning choruses for longer than 30 s and at a time when a chorus simultaneously emerged in a distant part of the forest.     

Patterns of Rarity in the Birds of the Atlantic Forest of Brazil

Patterns of rarity in species are generally explained by several factors: evolutionary history, spatial distribution, and genetic structure of each taxon. Human intervention also leads to or increases rarity in species. The discernment of causes of rarity is essential to the understanding of extinction patterns, and thus to devising conservation strategies. I examine patterns of rarity among bird species in the Atlantic forest region in Brazil, one of the most threatened ecosystems in the world. I assigned bird species to one of eight possible categories that differ in degree of vulnerability and that are based on three parameters of rarity: geographic distribution, habitat specificity, and population size. The Atlantic forest avifauna is a highly endangered group; 68% of the species are rare. Patterns of rarity among the birds in the region likely result from their specific ecologies or evolutionary histories. In addition, human alteration of natural habitats and hunting pressures have undoubtedly influenced rarity for a number of species.