JASs Reports Journal of Anthropological Sciences Vol. 83 (2005), pp. 89-109

The targeted : a re-evaluation of predation on New World

Bernardo Urbani

Department of Anthropology, University of Illinois at Urbana-Champaign, 109 Davenport Hall, 607 S Mathews Ave., Urbana, Illinois 61801, USA, e-mail: [email protected]

Summary – This work reviews the information related to predation on Neotropical primates by human and non-human predators. Paradoxically, humans have been systematically neglected while evaluating the potential effects of predation in the structure of non-human populations. Predation paradigms do not include humans in their propositions. In this review, it is shown that effectively humans are the main predators of monkey communities in the Neotropics. The results also suggest that humans do not fit with predation theoretical views given for non-human predators. Homo cultural hunting practices contribute to this situation. For example, humans seem to prefer larger primate groups that allow their location for hunting, or humans tend to prey primary larger monkeys with longer interbirth interval. In sum, it is suggested that since at least 11,000 years of human occupation in the Neotropics, Homo might have been played a fundamental role in the current organization and distribution of primate populations, including local extinctions. Humans seemed to have potentially influenced New World primate population in such short ecological time scale.

Keywords – Predatory behavior, ethnoprimatology, human hunting, predation paradigms, transdisciplinary perspective, conservation, Latin America.

Introduction works of the natural history of primates (e. g. Kinzey, 1997); however, reports of predation are “The indications are that man is the most extreme scarce. In part, this relates to difficulties serious enemy of howlers and that occasionally obtaining field data on predation because of its young may be attacked by ocelots” said low rate and rapid occurrence. In addition, Isbell Clarence Ray Carpenter in 1934 after observing (1994) suggested that limited predation monkeys in (Carpenter, observation might be related to the observers (field 1934: 129). In fact, it was his last conclusion in primatologists) that are normally not present the first systematic primate behavioral research when main predators are active (at night) and the conducted in the wild, and probably also the first presence of the observer may inhibit the scientific account of the impact of predators on appearance of predators during the day. feral primates. After this work, other researches However, predation and predation risk have have taken predation risk into account as a been considered important in modeling the potential factor influencing the evolution of structure and organization of the extant and sociality in general, and the social structure of extinct primate populations (see review: Isbell, primate populations (e.g. Cheney & Wragham, 1994). In this sense, it has been argued that some 1987). For example, since the 1960s chimpanzees “anti-predatory” behaviors such as vigilance, had been the subjects of the longest-term studies polyspecific associations, group cohesion, direct or even carried out for any wild ; but active defense and alarm calls are related to the paradoxically it was not until 1990, that the first selective pressure of predation on primates case of predation on chimpanzees by lions was (Cheney & Wragham, 1987; Isbell, 1994). Van reported (Tsukahara & Nishida 1990). In Schaik & Horstermann (1994) proposed a addition, predation has been systematically cited in hypothesis that suggested that the quantity of

the JASs is published by the Istituto Italiano di Antropologia www.isita-org.com 90 Predation on Neotropical Monkeys

primate males -and group composition- is not Recently, Miller (2002) added to the discussion only related to sexual competition but also to the concept of predation vulnerability in predation risk. Their hypothesis predicts that understanding primate social organization. She males are more vigilant than females, and play a defined vulnerability as the “qualitative measure of greater role in predator detection. the probability that an individual will be the victim In her literature review, Isbell (1994: 65-68) of a predator at any given moment” (Miller, 2002: characterized five patterns of predation on 2). She indicated that this fact may play a major role primates. These patterns are, 1) larger primates are in the foraging strategies and decision-making expected to be less vulnerable to predation than among primates. Miller (2002) suggested that are smaller primates; 2) an individual’s predation vulnerability might be evaluated using vulnerability to predation increased in unknown three different variables. A biological variable, in or unfamiliar areas (for a definition of which different characteristics that are “under vulnerability, see below: Miller, 2002); 3) primates genetic control” and expressed in the primate are more vulnerable in the upper canopy, in phenotype might be significant for a given anti- discontinuous forests and on the forest edges than predatory response (e. g. body size); a social variable in continuous, undisturbed forests; 4) predation is such as differences in rank, group size and group episodic and may be related to prey preferences composition; and an environmental variable such and different use of daily ranges and home ranges as the degree of predation vulnerability associated between predators and prey; and 5) terrestrial with foraging location and the quality of the cover primates have higher rates and risk of predation or refuge available to primates. than do arboreal primates. In addition, Chapman (2000) reviewed the predation avoidance Humans as primate predators hypothesis by examining patterns of group Isbell (1994) and Cheney & Wrangham structure and movement in 54 primate species. He (1987) briefly suggested that humans are the argued that living in groups, “[may] increased major predators on primates. Similarly, Boinski et probability of predator detection, … [create] al. (2000: 47) stated in their review, “primate greater confusion of a predator trying to focus on predators are restricted to nonhuman predators an individual prey, … [propitiate] a decreased because humans probably hunt most primates” probability of each individual being captured by (italics are mine). Thus, although the impact of predators, … and increased defense against humans as predators on primates has been noted, predators” (Chapman 2000: 27). few researchers have focused on the role of human However, Janson (1998) indicated that the predators may have played in non-human primate evaluation of predation should be done with social organization (see: Sponsel, 1997). Predation caution due to the fact that presumed anti- by humans has been under-emphasized in the predatory behaviors (e. g. alarm calls, vigilance, theoretical discussions about predation on non- living in larger groups) might be linked with other human primates. However, to hunt defined by the behaviors. For instance, he indicated that for some Oxford English Dictionary (Simpson & Weiner, primate species it might be beneficial to live in 1989: 496) is “the act of chasing wild animals for small groups that are difficult for predators to the purpose of catching or killing them”; not less detect. In addition, Hill & Dunbar (1998) than another form of predation. Consequently, as suggested that predation risk is a more indicated by Mittermeier (1987) and Chapman & comprehensive parameter for evaluating predation Peres (2001), New World primates are highly on primates than predation rate. They argued that threaten due to intense hunting or human potentially the role of predation as selective predation. Nevertheless, humans had often been pressure might be more important for primates neglected in the literature on predation in when the risk of predation is perceived. In primates (Sponsel 1997). In this paper I will addition, they indicated that predator-prey examine primate prey-human predator relationship should be evaluated as the potentiality relationship in the Neotropics. So, I address the that a given primate species might be recovered following questions, from predatory events by adjusting its behavior. a) Is there evidence that predators affect the B. Urbani 91

structure of primate communities in Neotropical human vertebrates that prey on Neotropical forests? primates. b) How human predators differ from non- b) Homo predator: refers only to Amerindian human predators in prey choice, hunting hunters of monkeys in the tropical forests. In this techniques and potential affect on primate social work only Amerindians groups were included, organization? considering the assumption that are the human b) Are humans primary agents that influence populations that preyed on monkeys inheriting ecological changes such as local extinctions of long-term traditional hunting techniques and primate populations? knowledge in the Neotropics (for Campesinos - Latin American creoles-, see discussion). Methods c) Prey: the targeted obtained by non-Homo or Homo predators. Information of predation on Neotropical Associated with this entry, in the cells of the primates was compiled (see Appendix). Only Appendix, information on the numbers of observed predation cases on monkeys by direct observed predation cases, age/sex of the prey, and sightings or from inspection of alimentary samples ranking of the prey is collected. In this sense, 1) such as primate skeletal material in predator nests Number of observed predation cases: refer to the and primate remains in predator feces were quantity of the proved cases of predation on included. For this purpose, a review of papers monkeys; 2) Age/Sex: age and sex of the prey; and related to predation and hunting on primates in 3) Ranking prey: only for the cases of Homo the Neotropics was systematically prepared from predators, refers to the rank for a given primate three bioanthropological -primatological- and genus compared to all mammalian preys. ethnographical sources. d) Total of prey: the sum of all preys for a given First, a search was conducted of the major predator and the number of prey in general. anthropological and biological databases such as e) Observation period: the total amount of time of Academic Search Elite, Anthropological Index, the primatological study or the time that the Anthropological Literature, Biological Abstracts ethnographer spent in the Amerindian (BIOSIS), Ecology Abstracts, JSTOR (including community in which the primate hunting events 35 ecological and anthropological journals dated were observed. from 1867 to 1999), PrimateLit and Zoological f) Frequency of predation: relationship between Record. the “Number of observed predation cases” per Secondly, a bibliographic compilation was hour. For both, primatological and ethnographical prepared from the reviews published by data, in which the observation period was not McDonald (1977), Coimbra-Filho & Mittermeier reported by hour, I used the following conversion: (1981), Beckerman & Sussenbach (1983), 1 day = 10 hours, 1 month = 15 days, 1 year = 12 Mittermeier (1987), Mittermeier et al. (1988), months. If the observation period was not Kinzey (1997), Sponsel (1997), Robinson & explicitly reported, it is included as unknown. Redford (1994), Boinski et al. (2000), Cowlishaw g) Locality: site in which the observation was & Dunbar (2000), Garber & Bicca-Marques done, including the region and the country. (2002), Di Fiore (2002), Fuentes & Wolfe (2002), Urbani (2002) and Cormier (2003). Third, the Results four main primatological journals American Journal of Primatology (1981-2003), Folia In Appendix, I summarizes the data archived Primatologica (1963-2003), International Journal from the bibliographical search on hunting and of Primatology (1980-2003), Primates (1957- predation on Neotropical monkeys. A total of 89 2003), and the ethnological Journal of entries were obtained, 33 (37.1% for non-Homo Ethnobiology (1981-2003) were also examined. predators and 56 (62.9%) for Homo predators. For From the compiled references, the following non-Homo predators, 51.5% the events were information was obtained (see the Appendix): recorded in non-Amazonian forests (17/33, incl. a) Non-Homo predator: corresponds with non- the Guianas, Central American and subtropic 92 Predation on Neotropical Monkeys

forests), and the other 48.5% (16/33) in the Amazonia. In the case of Homo predators, 69.6% (39/56) of the cases were in Amazonia while 30.4% (17/56) were in non-Amazonian forests. The frequency of predation was quite similar from 0.045/hour for non-Homo predators to 0.048/hour in Homo predators (Appendix). Among all non-Homo predators, avian predators appear to be the most common. Of 32 cases, harpy (Harpia harpija, 21.2%, 7/33) and crested (Morphnus guianensis, 12.1%, 4/33) are the most common predators. Among terrestrial non-Homo predators, (Panthera 12, p<0.01). Certainly, I am aware that some of onca) accounted for 15.2% (5/33) of the the human data collected for this comparison observations. Ten other non-Homo predators might imply the use of shotguns. For that reason, accounted for 51.5% (17/33) of predatory events. in order to model the potential hunting without For non-Homo predators it was not possible firearms, I reduced ten times the Homo predator determine the sex/age classes of the monkeys subtotals and compared it with the non-Homo preyed upon (69%, 29/42). However, for those predator dataset without modification. Hence, predation cases in which the information was after this subtraction and using the same statistical available, 21.4% (9/42) represent young animals test, the results are still highly different. (pooling together infants, juveniles and subadults) A ranking of New World monkeys arranged and the remaining 9.5% (4/42) were adults, while by the number of individuals hunted is presented 69.1% were of unknown age. In the few cases in in Tab. 2. The life history data, -mean body mass which the sex was observed, 9.5% (4/42) were and interbirth interval-, were obtained from males whereas 7.1% (3/42) females, and the rest Kappeler & Pereira (2003). As showed in Tab. 2, 83.4% remained of unknown sex. For Homo Cebus is the most commonly hunted monkey for predators, in all cases the specific sex/age classes both Homo and non-Homo predators. For non- were unknown. In addition, as indicated in the Homo predators, small and medium sized primates discussion (see below) 53.6% (30/56) of the are selected more frequently. Among humans, entries of Homo predators explicitly stated that large bodied genera such as Lagothrix, Ateles and these human groups hunt at least one of the non- Alouatta are the preferred prey. Therefore, for Homo predators listed in Appendix. Moreover, for Homo predators, a larger primate body mass plays Homo predators it is important to identify the a major role in prey choice. ranking of monkey preferences among all mammal prey. It is interesting to note that among Discussion Appendix entries, monkeys are among the five most common preys (27.8%, 42/151; see Tab. 1), Humans first arrived to lowland South the rest 72.5% are for primates preferred over the America around 11,000 years ago (Amazonia: six position of preference (15.9%, 24/151) or it is Roosevelt et al., 1996). The technology they used unknown the choice rank among Amerindian for hunting included primarily stone spear points groups (56.3%, 85/151). as could be recovered from Pleistocene I tested the prediction that Homo predators archaeological sites (Roosevelt et al., 1996). Thus, have higher proficiency obtaining New World humans have likely played a fundamental role in monkeys than non-Homo predators. Proficiency hunting different prey since that time. was defined as the number of successful hunts of a Currently, based on this review of predation on given primate prey. Both, Homo predators and New World non-human primates, it appears that non-Homo predators were compared using G-test. Homo is the main predator of them when The results indicate that humans are more compared with non-Homo predators. In a study of proficient predators of New World primates (df= protein requirements for Amazonian Amerindian B. Urbani 93

population, it was suggested that a daily diet population in the Brazilian should have at least 40-50 g/day per capita of Amazon (Peres, 1991; see below). proteins where game meat and fish occupied an On the other hand, humans do not fit well in important place apart of plant proteins from other proposed predatory predictions of predator- sources such as mandioca (Gross, 1975, for primate prey interactions. In this sense, various discussion see: Beckerman, 1979). For obtaining anti-predatory behaviors such as mobbing, active Neotropical animal game, hunting practices are defense (e. g. throwing branches) and alarm calls culturally based (Beckerman & Sussenbach, 1983; are not successful for the monkeys because they Sponsel, 1997; Lizot, 1979; Lizarralde, 2002). For would be easily detected and killed by humans. example, Lizot (1979) indicated that food For example, some Amerindian groups such as the preferences and taboos among Yanomami groups Shirián of the upper Paragua River, , determine the selection and consumption of performed monkey calls in order to wait for an particular food items, particularly key game acoustical response to locate their potential prey animals. Also, cultural practices might influence (Urbani, pers. obs.). On the other hand, in the inhibition for eating some monkeys. Among , the Lacandón people hear the roaring of the Desana of Amazonian , monkeys are the monkeys from a long distance to chase and prohibited meat for children, while howler hunt them (Baer & Merrifield, 1972). Moreover, monkeys are considered evil omens (Reichel- other so-called anti-predatory behaviors like Dolmatoff, 1971). In addition, humans are polyspecific associations and intragroup distance dramatically decreasing the non-human predator might aid human hunters in locating the primate populations. For instance, over-hunting in the groups. For instance, Balée (1985) reported that Neotropics has created the so-called “empty hunting howler monkeys by the Ka’apor in , forests,” were major were removed from take them an average of just 50 minutes from the their natural habitats (Redford, 1992; Robinson, moment they left from and returned to the 1993). The result is that in some tropical areas, settlement after the hunting party. This is the humans maybe are the only threat to feral monkey smallest amount of time used for hunting any populations (Cowlishaw & Dunbar, 2000). This mammalian game for this Amerindian group, the has been documented in terms of the decreasing next nearest are collared peccaries that required in 94 Predation on Neotropical Monkeys

average 240 minutes, so, almost five times more be more easily found by hunters. For instance, than howlers. practically entire groups of woolly monkeys (130 The prediction that smaller-bodied size individuals) and 11 howler monkeys were killed primates are more vulnerable to predators than in one day by a Siona-Secoya hunting party larger bodied primates does not fit for human composed by 286 persons in the Ecuadorian predators. It has been observed that large Amazon (Vickers, 1980). The idea that primate atelines are the preferred prey for many human groups might create predator confusions and groups, and are systematically selected as thereby decrease the feasibility that each hunting targets (Mena et al., 2000; Alvard & individual may be preyed while increasing the Kaplan, 1991). Another prediction suggested defense against the predators, does not appears that the vulnerability to predation might be to function as a potential anti-predatory higher in unfamiliar areas for the primates. Local adaptation for human hunters. Hunters may kill human populations tend to know the monkeys the most of the whole group or just select home range, and in some cases hunters may particular individuals based on cultural predict the movements of primates populations preferences. In this sense, female primates may through their knowledge of fruiting patterns of be selected to hunt in order to obtain offspring as particular key tree species eaten by monkeys or pets. In the case of Ateles sp., females may be identifying animal feces in the forest (e. g. the selected because they are considered more “tasty” Makuna: Århem, 1976). On the other hand, the than males (Waimiri-Atroari: Souza-Mazurek et prediction that primates are more vulnerable in al., 2000), more “tasty” than other monkey species the canopy and forest edges may apply when being such as Saimiri oerstedii (Guaymi: González- hunted by human predators. Based on interviews Kirchner & Sainz de la Maza, 1998), or even are in the Venezuelan Guayana and eastern Venezuela, considered “better” hunting games during the I received information from hunters (Amerindians rainy season because this monkey species is fatter and Creoles) indicating that hunting proficiency is during this tree fruiting period (Matsigenka: higher when monkeys are exposed in the forest Shepard, 2002). canopies rather than in dense tangles. They Boinski & Chapman (1995) provided new indicated that it is easier to hunt a insights on potential directions for testing than tend to be found in the highest canopy than hypotheses of predation on primates. They a sometimes found in tangle argued that comparisons of predation rates with understory forests strata or secondary forests. In group size might represent a bias because of large this sense, Carneiro (1970) indicated that in intraspecific variability in primate group size. Peruvian Amahuaca hunting, success decreases However, they suggest that other standards for with higher dense foliage; however, he reported comparisons like prey interbirth intervals might that this Amerindian group practiced tree be more fruitful for understanding the effects of climbing for hunting howler monkeys. predation on primates. It is important to note The idea that predation is episodic must be that despite the low frequencies of predation for re-evaluated when dealing with human hunters. both predators, primates with longer interbirth Homo hunts primates on a regular basis and one intervals such as Ateles sp. and Lagothrix sp. are hunting -predatory- episode by humans might the preferred targets among human hunters (e. result in the killing of many members of the g. Yanomamö: Saffirio & Scaglion, 1982; monkey group. For example, Lizarralde (2002) Matsigenka: Shepard, 2002; Tabl. 1). This may reported the case of a single hunting day in which result in low levels of primate population five Barí men returned to the village with 16 recovery and also local extinctions. spider monkeys (Ateles hybridus). Boinski & Chapman (1995: 2) state “on an The consequences of living in group for evolutionary time scale, increased predation avoiding predation as reviewed by Chapman pressure may favor large groups, but on a shorter (2000) must be revisited for human predators. ecological time scale, high predation levels may Contrary to the prediction, living in a large decrease group size directly, simply through the group is a disadvantage, because monkeys may death of animals.” In this regard, they indicated B. Urbani 95

that G. B. Stanford studies of chimpanzees (Pan extinctions of faunal populations and probably - troglodytes) predation on red colobus monkeys still unknown- contributing to shape the structure (Procolobus badius) is an instructive example of the of current primate populations as might be effect of predation on an ecological time scale. tentatively inferred from the results. There is Chimpanzees hunting on red colobus (76 current evidence to support this idea. For instance, observed cases in four years) have reduced to Souza-Mazurek et al. (2000 579; 591) indicated almost the half the size of the monkey population that among the Waimiri-Atroari in northern that inhabit the chimpanzee groups’ range Brazil, “sex ratios of spider monkeys [Ateles compared to the red colobus populations that live paniscus] killed were heavily biased towards outside this range (e.g. Gombe site: Stanford, females indicating a stronger hunting pressure on 1998 vs Kibale site: Struhsaker, 1975). In other those individuals.” Thus, it seems that potential words, in the field sites in which Pan density is sex-based differences in hunting selection are higher, Procolobus density is lower (Stanford, affecting the group structure of feral Neotropical 1998). primates groups; naturally more detailed studies A similar effect may be occurring in the to evaluate the effect of selective hunting on the Neotropics between human and some monkey primate population dynamic are needed. populations. Since the Prehispanic period, Moreover, Peres (1991) described local Amerindians established a close relationship with extinctions of woolly monkey (Lagothrix sp.) non-human primates from cosmological believes populations as a consequence of intense hunting to the thought of monkeys as food (Baker, 1992,; in the Amazonia. He said that woolly monkey van Akkeren, 1998; Karadimas, 1999, Braakhuis, population density (number/km2) varied from 0 1998, García del Cueto, 1989; Urbani & Gil, and 7 in hunted sites to 17 and 30 in non- 2001; Cormier, 2003). In the present work only hunted sites. In addition, Creole populations in Amerindians groups were considered, assuming Latin America intensively hunt primates for that are the human populations that first entered market networks that expand the limits of hunting the New World while using long-term traditional from family consumption to hunting for hunting techniques. The predecessors of current commercial purposes (Mittermeier, 1987, for Amerindian populations occupied the tropical understanding Amazonian Caboblos -Brazilian Americas since circa 11,000 year ago to Creoles- economy: Nugent, 1993). Bush meat is a approximately 1,250-1,600 A. D., intensely using preferred food source in the local marts. For areas like lowland Panama and Brazil that have example, Castro et al. (1975) found that in six been historically considered “pristine” months, in the popular market of Iquitos, , in the New World (Bush & Colinvaux, 1990; were sold 1,700 Lagothrix sp., 1,396 Cebus sp., Colinvaux & Bush, 1991; Heckenberger et al., 557 Alouatta sp., 321 Pithecia sp. and 198 Ateles 2003). In addition, there is direct evidence from sp. (approximate calculation by B. U.). Thus, an archaeological site in northern Venezuela that together with the Congo Basin in Africa where the reflected potential consumption or used for other bush meat crisis is aggressively threatening primate purposes (e. g. pets) of red howler monkeys from populations (Peterson, 2003), Amazonia appears at least 3,000 years B. P. (Urbani & Gil, 2001). In to be the other geographical area critically threaten principle, these human groups may have by hunting (Mittermeier, 1987; see Results). influenced the distribution/survivorship of current Then, to the question, why New World primate populations, as might be the case of primates adopt the above indicated anti- potential former populations of white-faced predatory behaviors even if they are not efficient capuchin monkeys (Cebus capucinus) in the against their main predator, Homo sapiens?. Two northern Mayan region (Baker, 1992). In dimensions might be playing a role on it. First, addition, MacPhee & Horovitz (2002) suggested humans have interacted with New World that probably the Pleistocene Antillean monkey monkeys during just a short timeframe (~11,000 Xenothrix mcgregori might be extinct due to years) compared to the long-time period since human influence. So, intense hunting pressures primates had been found in the New World; the over 10,000 years might directly influence local late Oligocene when Branisella boliviana 96 Predation on Neotropical Monkeys

inhabited the central part of (including primatologists), in order to understand (Kay et al. 2002). On the other hand, human potential differences in primate social structure social practices and material culture (particularly due to human presence or impact. weapons) take part in the equation of predation of Neotropical monkeys, being factors unique to Homo in the Neotropics. Nevertheless, extensive Acknowledgments field research is needed to properly address this issue. Many thanks to Loretta A. Cormier, Manuel Probably one of the best ways to inquiry Lizarralde, Leslie E. Sponsel and Robert S. Voss for about the relationship between Homo and New sending me their publications for a previous World primates, including in particular hunting bibliographic review (Urbani, 2002) that were -predatory- practices, is to look into new useful for thinking in this work. To the UIUC insights from ethnoprimatological perspectives library staff for make available valuable help in on the conceptualization of monkeys by references searching. To Paul Garber and the Amerindians; a research line that have been just anonymous referees for their critical suggestions. To initiated after Sponsel (1997) (in the Tania for her passion in the conservation of tropical Neotropics: Fleck et al., 1999; Cormier, 2002, rainforests and her enormous love. B. Urbani is 2003; Lizarralde, 2002; Shepard, 2002; Urbani granted by a Fulbright-OAS Scholarship. & Gil, 2001). Many Amerindian groups have special tied relations with monkeys, and associated and classified them as “beings like References humans” (e.g. Kalapalo: Basso, 1973). For instance, among the Mekranoti, a Tupi language Alvard M. 1993. Testing the “Ecologically noble Amerindian group related to the Kayapo group, savage” hypothesis: Interpecific prey choice indicated that other Amerindian groups used the by Piro hunters of Amazonian Perú. Hum. word “Kaya-po” for referring them; the word Ecol., 21: 355-387. “Kayapo” means the people that “resembled Alvard M. 1995. Intraspecific prey choice by monkeys” (Werner, 1985: 173). Ethnoprimatological Amazonian hunters. Curr. Anthropol., 36: studies are fundamental for understanding the 789-818. relationship between human and non-human Alvard M. 1995. Shotguns and sustainable primates. Moreover, these works will be essential hunting in the neotropics. Oryx, 29: 58-66. for understanding the past and present inter- Alvard M. & Kaplan H. 1991. Procurement connection of both sympatric primates and even technology and prey mortality among more, to comprehend the still unknown but indigenous neotropical hunters. In M.C. critical future of them. In addition, at the Stiner (ed): Human predators and prey moment there are few systematic mentions mortality, pp. 79-104. Westview Press, describing the New Word primate reactions to Boulder. human primates, and in principle there are Århem K. 1976. Fishing and hunting among the practically no works specifically describing how Makuma: Economy, ideology and ecological humans have potentially affected social and adaptation in the northwest Amazon. Ann. grouping behaviors in Neotropical monkeys (for Report of the Gotenborgs Etnografiska Museum, Old World primates: Coppinger & Maguire, : 27-44. 1980; Bshary, 2001; Tenaza, 1990; Tenaza & Baer P. & Merrifield W.R. 1972. Los Lacandones de Tilson, 1985). This is an interesting area for future México. Dos Estudios. Instituto Nacional primatological research in the tropical Americas. Indigenista, Secretaría de Educación Pública, Moreover, recent data on naïve behaviors and Mexico City. group composition of wild primates (for Baker M. 1992. Capuchin monkeys (Cebus chimpanzees: Morgan & Sanz, 2003) may be capucinus) and the Ancient Maya. Ancient fundamental for comparing with primate Mesoam., 3: 219-228. populations subjected to often human contact Baleé W. 1985. Ka’apor ritual hunting. Hum. B. Urbani 97

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