the targeted monkey: a re-evaluation of predation on new world primates vol83/urbani.pdf ·...

JASs ReportsJournal of Anthropological Sciences

Vol. 83 (2005), pp. 89-109

the JASs is published by the Istituto Italiano di Antropologia www.isita-org.com

The targeted monkey: a re-evaluation of predation onNew World primates

Bernardo Urbani

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

Summary – This work reviews the information related to predation on Neotropical primates by humanand non-human predators. Paradoxically, humans have been systematically neglected while evaluating thepotential effects of predation in the structure of non-human primate populations. Predation paradigms do notinclude humans in their propositions. In this review, it is shown that effectively humans are the main predatorsof monkey communities in the Neotropics. The results also suggest that humans do not fit with predationtheoretical views given for non-human predators. Homo cultural hunting practices contribute to thissituation. For example, humans seem to prefer larger primate groups that allow their location for hunting, orhumans tend to prey primary larger monkeys with longer interbirth interval. In sum, it is suggested that sinceat least 11,000 years of human occupation in the Neotropics, Homo might have been played a fundamentalrole in the current organization and distribution of primate populations, including local extinctions. Humansseemed 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

“The indications are that man is the mostserious enemy of howlers and that occasionallyyoung animals may be attacked by ocelots” saidClarence Ray Carpenter in 1934 after observingmantled howler monkeys in Panama (Carpenter,1934: 129). In fact, it was his last conclusion inthe first systematic primate behavioral researchconducted in the wild, and probably also the firstscientific account of the impact of predators onferal primates. After this work, other researcheshave taken predation risk into account as apotential factor influencing the evolution ofsociality in general, and the social structure ofprimate populations (e.g. Cheney & Wragham,1987). For example, since the 1960s chimpanzeeshad been the subjects of the longest-term studieseven carried out for any wild mammal; butparadoxically it was not until 1990, that the firstcase of predation on chimpanzees by lions wasreported (Tsukahara & Nishida 1990). Inaddition, predation has been systematically cited in

works of the natural history of primates (e. g.Kinzey, 1997); however, reports of predation areextreme scarce. In part, this relates to difficultiesobtaining field data on predation because of itslow rate and rapid occurrence. In addition, Isbell(1994) suggested that limited predationobservation might be related to the observers (fieldprimatologists) that are normally not presentwhen main predators are active (at night) and thepresence of the observer may inhibit theappearance of predators during the day.

However, predation and predation risk havebeen considered important in modeling thestructure and organization of the extant andextinct primate populations (see review: Isbell,1994). In this sense, it has been argued that some“anti-predatory” behaviors such as vigilance,polyspecific associations, group cohesion, direct oractive defense and alarm calls are related to theselective pressure of predation on primates(Cheney & Wragham, 1987; Isbell, 1994). VanSchaik & Horstermann (1994) proposed ahypothesis that suggested that the quantity of

primate males -and group composition- is notonly related to sexual competition but also topredation risk. Their hypothesis predicts thatmales are more vigilant than females, and play agreater role in predator detection.

In her literature review, Isbell (1994: 65-68)characterized five patterns of predation onprimates. These patterns are, 1) larger primates areexpected to be less vulnerable to predation thanare smaller primates; 2) an individual’svulnerability to predation increased in unknownor unfamiliar areas (for a definition ofvulnerability, see below: Miller, 2002); 3) primatesare more vulnerable in the upper canopy, indiscontinuous forests and on the forest edges thanin continuous, undisturbed forests; 4) predation isepisodic and may be related to prey preferencesand different use of daily ranges and home rangesbetween predators and prey; and 5) terrestrialprimates have higher rates and risk of predationthan do arboreal primates. In addition, Chapman(2000) reviewed the predation avoidancehypothesis by examining patterns of groupstructure and movement in 54 primate species. Heargued that living in groups, “[may] increasedprobability of predator detection, … [create]greater confusion of a predator trying to focus onan individual prey, … [propitiate] a decreasedprobability of each individual being captured bypredators, … and increased defense againstpredators” (Chapman 2000: 27).

However, Janson (1998) indicated that theevaluation of predation should be done withcaution due to the fact that presumed anti-predatory behaviors (e. g. alarm calls, vigilance,living in larger groups) might be linked with otherbehaviors. For instance, he indicated that for someprimate species it might be beneficial to live insmall groups that are difficult for predators todetect. In addition, Hill & Dunbar (1998)suggested that predation risk is a morecomprehensive parameter for evaluating predationon primates than predation rate. They argued thatpotentially the role of predation as selectivepressure might be more important for primateswhen the risk of predation is perceived. Inaddition, they indicated that predator-preyrelationship should be evaluated as the potentialitythat a given primate species might be recoveredfrom predatory events by adjusting its behavior.

Recently, Miller (2002) added to the discussionthe concept of predation vulnerability inunderstanding primate social organization. Shedefined vulnerability as the “qualitative measure ofthe probability that an individual will be the victimof a predator at any given moment” (Miller, 2002:2). She indicated that this fact may play a major rolein the foraging strategies and decision-makingamong primates. Miller (2002) suggested thatpredation vulnerability might be evaluated usingthree different variables. A biological variable, inwhich different characteristics that are “undergenetic control” and expressed in the primatephenotype might be significant for a given anti-predatory response (e. g. body size); a social variablesuch as differences in rank, group size and groupcomposition; and an environmental variable suchas the degree of predation vulnerability associatedwith foraging location and the quality of the coveror refuge available to primates.

Humans as primate predatorsIsbell (1994) and Cheney & Wrangham

(1987) briefly suggested that humans are themajor predators on primates. Similarly, Boinski etal. (2000: 47) stated in their review, “primatepredators are restricted to nonhuman predatorsbecause humans probably hunt most primates”(italics are mine). Thus, although the impact ofhumans as predators on primates has been noted,few researchers have focused on the role of humanpredators may have played in non-human primatesocial organization (see: Sponsel, 1997). Predationby humans has been under-emphasized in thetheoretical discussions about predation on non-human primates. However, to hunt defined by theOxford English Dictionary (Simpson & Weiner,1989: 496) is “the act of chasing wild animals forthe purpose of catching or killing them”; not lessthan another form of predation. Consequently, asindicated by Mittermeier (1987) and Chapman &Peres (2001), New World primates are highlythreaten due to intense hunting or humanpredation. Nevertheless, humans had often beenneglected in the literature on predation inprimates (Sponsel 1997). In this paper I willexamine primate prey-human predatorrelationship in the Neotropics. So, I address thefollowing questions,

a) Is there evidence that predators affect the

90 Predation on Neotropical Monkeys

structure of primate communities in Neotropicalforests?

b) How human predators differ from non-human predators in prey choice, huntingtechniques and potential affect on primate socialorganization?

b) Are humans primary agents that influenceecological changes such as local extinctions ofprimate populations?

Methods

Information of predation on Neotropicalprimates was compiled (see Appendix). Onlyobserved predation cases on monkeys by directsightings or from inspection of alimentary samplessuch as primate skeletal material in predator nestsand primate remains in predator feces wereincluded. For this purpose, a review of papersrelated to predation and hunting on primates inthe Neotropics was systematically prepared fromthree bioanthropological -primatological- andethnographical sources.

First, a search was conducted of the majoranthropological and biological databases such asAcademic Search Elite, Anthropological Index,Anthropological Literature, Biological Abstracts(BIOSIS), Ecology Abstracts, JSTOR (including35 ecological and anthropological journals datedfrom 1867 to 1999), PrimateLit and ZoologicalRecord.

Secondly, a bibliographic compilation wasprepared from the reviews published byMcDonald (1977), Coimbra-Filho & Mittermeier(1981), Beckerman & Sussenbach (1983),Mittermeier (1987), Mittermeier et al. (1988),Kinzey (1997), Sponsel (1997), Robinson &Redford (1994), Boinski et al. (2000), Cowlishaw& Dunbar (2000), Garber & Bicca-Marques(2002), Di Fiore (2002), Fuentes & Wolfe (2002),Urbani (2002) and Cormier (2003). Third, thefour main primatological journals AmericanJournal of Primatology (1981-2003), FoliaPrimatologica (1963-2003), International Journalof Primatology (1980-2003), Primates (1957-2003), and the ethnological Journal ofEthnobiology (1981-2003) were also examined.

From the compiled references, the followinginformation was obtained (see the Appendix):a) Non-Homo predator: corresponds with non-

human vertebrates that prey on Neotropicalprimates. b) Homo predator: refers only to Amerindianhunters of monkeys in the tropical forests. In thiswork only Amerindians groups were included,considering the assumption that are the humanpopulations that preyed on monkeys inheritinglong-term traditional hunting techniques andknowledge in the Neotropics (for Campesinos -Latin American creoles-, see discussion).c) Prey: the targeted New World monkey genusobtained by non-Homo or Homo predators.Associated with this entry, in the cells of theAppendix, information on the numbers ofobserved predation cases, age/sex of the prey, andranking of the prey is collected. In this sense, 1)Number of observed predation cases: refer to thequantity of the proved cases of predation onmonkeys; 2) Age/Sex: age and sex of the prey; and3) Ranking prey: only for the cases of Homopredators, refers to the rank for a given primategenus compared to all mammalian preys.d) Total of prey: the sum of all preys for a givenpredator and the number of prey in general.e) Observation period: the total amount of time ofthe primatological study or the time that theethnographer spent in the Amerindiancommunity in which the primate hunting eventswere observed. f ) Frequency of predation: relationship betweenthe “Number of observed predation cases” perhour. For both, primatological and ethnographicaldata, in which the observation period was notreported by hour, I used the following conversion:1 day = 10 hours, 1 month = 15 days, 1 year = 12months. If the observation period was notexplicitly reported, it is included as unknown. g) Locality: site in which the observation wasdone, including the region and the country.

Results

In Appendix, I summarizes the data archivedfrom the bibliographical search on hunting andpredation on Neotropical monkeys. A total of 89entries were obtained, 33 (37.1% for non-Homopredators and 56 (62.9%) for Homo predators. Fornon-Homo predators, 51.5% the events wererecorded in non-Amazonian forests (17/33, incl.the Guianas, Central American and subtropic

B. Urbani 91

forests), and the other 48.5% (16/33) in theAmazonia. In the case of Homo predators, 69.6%(39/56) of the cases were in Amazonia while30.4% (17/56) were in non-Amazonian forests.The frequency of predation was quite similar from0.045/hour for non-Homo predators to0.048/hour in Homo predators (Appendix).

Among all non-Homo predators, avianpredators appear to be the most common. Of 32cases, harpy eagles (Harpia harpija, 21.2%, 7/33)and crested eagle (Morphnus guianensis, 12.1%,4/33) are the most common predators. Amongterrestrial non-Homo predators, jaguars (Pantheraonca) accounted for 15.2% (5/33) of theobservations. Ten other non-Homo predatorsaccounted for 51.5% (17/33) of predatory events.For non-Homo predators it was not possibledetermine the sex/age classes of the monkeyspreyed upon (69%, 29/42). However, for thosepredation cases in which the information wasavailable, 21.4% (9/42) represent young animals(pooling together infants, juveniles and subadults)and the remaining 9.5% (4/42) were adults, while69.1% were of unknown age. In the few cases inwhich the sex was observed, 9.5% (4/42) weremales whereas 7.1% (3/42) females, and the rest83.4% remained of unknown sex. For Homopredators, in all cases the specific sex/age classeswere unknown. In addition, as indicated in thediscussion (see below) 53.6% (30/56) of theentries of Homo predators explicitly stated thatthese human groups hunt at least one of the non-Homo predators listed in Appendix. Moreover, forHomo predators it is important to identify theranking of monkey preferences among allmammal prey. It is interesting to note that amongAppendix entries, monkeys are among the fivemost common preys (27.8%, 42/151; see Tab. 1),the rest 72.5% are for primates preferred over thesix position of preference (15.9%, 24/151) or it isunknown the choice rank among Amerindiangroups (56.3%, 85/151).

I tested the prediction that Homo predatorshave higher proficiency obtaining New Worldmonkeys than non-Homo predators. Proficiencywas defined as the number of successful hunts of agiven primate prey. Both, Homo predators andnon-Homo predators were compared using G-test.The results indicate that humans are moreproficient predators of New World primates (df=

12, p<0.01). Certainly, I am aware that some ofthe human data collected for this comparisonmight imply the use of shotguns. For that reason,in order to model the potential hunting withoutfirearms, I reduced ten times the Homo predatorsubtotals and compared it with the non-Homopredator dataset without modification. Hence,after this subtraction and using the same statisticaltest, the results are still highly different.

A ranking of New World monkeys arrangedby the number of individuals hunted is presentedin Tab. 2. The life history data, -mean body massand interbirth interval-, were obtained fromKappeler & Pereira (2003). As showed in Tab. 2,Cebus is the most commonly hunted monkey forboth Homo and non-Homo predators. For non-Homo predators, small and medium sized primatesare selected more frequently. Among humans,large bodied genera such as Lagothrix, Ateles andAlouatta are the preferred prey. Therefore, forHomo predators, a larger primate body mass playsa major role in prey choice.

Discussion

Humans first arrived to lowland SouthAmerica around 11,000 years ago (Amazonia:Roosevelt et al., 1996). The technology they usedfor hunting included primarily stone spear pointsas could be recovered from Pleistocenearchaeological sites (Roosevelt et al., 1996). Thus,humans have likely played a fundamental role inhunting different animal prey since that time.Currently, based on this review of predation onNew World non-human primates, it appears thatHomo is the main predator of them whencompared with non-Homo predators. In a study ofprotein requirements for Amazonian Amerindian


population, it was suggested that a daily dietshould have at least 40-50 g/day per capita ofproteins where game meat and fish occupied animportant place apart of plant proteins fromsources such as mandioca (Gross, 1975, fordiscussion see: Beckerman, 1979). For obtainingNeotropical animal game, hunting practices areculturally based (Beckerman & Sussenbach, 1983;Sponsel, 1997; Lizot, 1979; Lizarralde, 2002). Forexample, Lizot (1979) indicated that foodpreferences and taboos among Yanomami groupsdetermine the selection and consumption ofparticular food items, particularly key gameanimals. Also, cultural practices might influencethe inhibition for eating some monkeys. Amongthe Desana of Amazonian Colombia, monkeys areprohibited meat for children, while howlermonkeys are considered evil omens (Reichel-Dolmatoff, 1971). In addition, humans aredramatically decreasing the non-human predatorpopulations. For instance, over-hunting in theNeotropics has created the so-called “emptyforests,” were major mammals were removed fromtheir natural habitats (Redford, 1992; Robinson,1993). The result is that in some tropical areas,humans maybe are the only threat to feral monkeypopulations (Cowlishaw & Dunbar, 2000). Thishas been documented in terms of the decreasing

woolly monkey population in the BrazilianAmazon (Peres, 1991; see below).

On the other hand, humans do not fit well inother proposed predatory predictions of predator-primate prey interactions. In this sense, variousanti-predatory behaviors such as mobbing, activedefense (e. g. throwing branches) and alarm callsare not successful for the monkeys because theywould be easily detected and killed by humans.For example, some Amerindian groups such as theShirián of the upper Paragua River, Venezuela,performed monkey calls in order to wait for anacoustical response to locate their potential prey(Urbani, pers. obs.). On the other hand, inMexico, the Lacandón people hear the roaring ofthe monkeys from a long distance to chase andhunt them (Baer & Merrifield, 1972). Moreover,other so-called anti-predatory behaviors likepolyspecific associations and intragroup distancemight aid human hunters in locating the primategroups. For instance, Balée (1985) reported thathunting howler monkeys by the Ka’apor in Brazil,take them an average of just 50 minutes from themoment they left from and returned to thesettlement after the hunting party. This is thesmallest amount of time used for hunting anymammalian game for this Amerindian group, thenext nearest are collared peccaries that required in

B. Urbani 93


average 240 minutes, so, almost five times morethan howlers.

The prediction that smaller-bodied sizeprimates are more vulnerable to predators thanlarger bodied primates does not fit for humanpredators. It has been observed that largeatelines are the preferred prey for many humangroups, and are systematically selected ashunting targets (Mena et al., 2000; Alvard &Kaplan, 1991). Another prediction suggestedthat the vulnerability to predation might behigher in unfamiliar areas for the primates. Localhuman populations tend to know the monkeyshome range, and in some cases hunters maypredict the movements of primates populationsthrough their knowledge of fruiting patterns ofparticular key tree species eaten by monkeys oridentifying animal feces in the forest (e. g. theMakuna: Århem, 1976). On the other hand, theprediction that primates are more vulnerable inthe canopy and forest edges may apply when beinghunted by human predators. Based on interviewsin the Venezuelan Guayana and eastern Venezuela,I received information from hunters (Amerindiansand Creoles) indicating that hunting proficiency ishigher when monkeys are exposed in the forestcanopies rather than in dense tangles. Theyindicated that it is easier to hunt a spider monkeythan tend to be found in the highest canopy thana capuchin monkey sometimes found in tangleunderstory forests strata or secondary forests. Inthis sense, Carneiro (1970) indicated that inPeruvian Amahuaca hunting, success decreaseswith higher dense foliage; however, he reportedthat this Amerindian group practiced treeclimbing for hunting howler monkeys.

The idea that predation is episodic must bere-evaluated when dealing with human hunters.Homo hunts primates on a regular basis and onehunting -predatory- episode by humans mightresult in the killing of many members of themonkey group. For example, Lizarralde (2002)reported the case of a single hunting day in whichfive Barí men returned to the village with 16spider monkeys (Ateles hybridus).

The consequences of living in group foravoiding predation as reviewed by Chapman(2000) must be revisited for human predators.Contrary to the prediction, living in a largegroup is a disadvantage, because monkeys may

be more easily found by hunters. For instance,practically entire groups of woolly monkeys (130individuals) and 11 howler monkeys were killedin one day by a Siona-Secoya hunting partycomposed by 286 persons in the EcuadorianAmazon (Vickers, 1980). The idea that primategroups might create predator confusions andthereby decrease the feasibility that eachindividual may be preyed while increasing thedefense against the predators, does not appearsto function as a potential anti-predatoryadaptation for human hunters. Hunters may killthe most of the whole group or just selectparticular individuals based on culturalpreferences. In this sense, female primates maybe selected to hunt in order to obtain offspring aspets. In the case of Ateles sp., females may beselected because they are considered more “tasty”than males (Waimiri-Atroari: Souza-Mazurek etal., 2000), more “tasty” than other monkey speciessuch as Saimiri oerstedii (Guaymi: González-Kirchner & Sainz de la Maza, 1998), or even areconsidered “better” hunting games during therainy season because this monkey species is fatterduring this tree fruiting period (Matsigenka:Shepard, 2002).

Boinski & Chapman (1995) provided newinsights on potential directions for testinghypotheses of predation on primates. Theyargued that comparisons of predation rates withgroup size might represent a bias because of largeintraspecific variability in primate group size.However, they suggest that other standards forcomparisons like prey interbirth intervals mightbe more fruitful for understanding the effects ofpredation on primates. It is important to notethat despite the low frequencies of predation forboth predators, primates with longer interbirthintervals such as Ateles sp. and Lagothrix sp. arethe preferred targets among human hunters (e.g. Yanomamö: Saffirio & Scaglion, 1982;Matsigenka: Shepard, 2002; Tabl. 1). This mayresult in low levels of primate populationrecovery and also local extinctions.

Boinski & Chapman (1995: 2) state “on anevolutionary time scale, increased predationpressure may favor large groups, but on a shorterecological time scale, high predation levels maydecrease group size directly, simply through thedeath of animals.” In this regard, they indicated

that G. B. Stanford studies of chimpanzees (Pantroglodytes) predation on red colobus monkeys(Procolobus badius) is an instructive example of theeffect of predation on an ecological time scale.Chimpanzees hunting on red colobus (76observed cases in four years) have reduced toalmost the half the size of the monkey populationthat inhabit the chimpanzee groups’ rangecompared to the red colobus populations that liveoutside this range (e.g. Gombe site: Stanford,1998 vs Kibale site: Struhsaker, 1975). In otherwords, in the field sites in which Pan density ishigher, Procolobus density is lower (Stanford,1998).

A similar effect may be occurring in theNeotropics between human and some monkeypopulations. Since the Prehispanic period,Amerindians established a close relationship withnon-human primates from cosmological believesto the thought of monkeys as food (Baker, 1992,;van Akkeren, 1998; Karadimas, 1999, Braakhuis,1998, García del Cueto, 1989; Urbani & Gil,2001; Cormier, 2003). In the present work onlyAmerindians groups were considered, assumingthat are the human populations that first enteredthe New World while using long-term traditionalhunting techniques. The predecessors of currentAmerindian populations occupied the tropicalAmericas since circa 11,000 year ago toapproximately 1,250-1,600 A. D., intensely usingareas like lowland Panama and Brazil that havebeen historically considered “pristine” rainforestsin the New World (Bush & Colinvaux, 1990;Colinvaux & Bush, 1991; Heckenberger et al.,2003). In addition, there is direct evidence froman archaeological site in northern Venezuela thatreflected potential consumption or used for otherpurposes (e. g. pets) of red howler monkeys fromat least 3,000 years B. P. (Urbani & Gil, 2001). Inprinciple, these human groups may haveinfluenced the distribution/survivorship of currentprimate populations, as might be the case ofpotential former populations of white-facedcapuchin monkeys (Cebus capucinus) in thenorthern Mayan region (Baker, 1992). Inaddition, MacPhee & Horovitz (2002) suggestedthat probably the Pleistocene Antillean monkeyXenothrix mcgregori might be extinct due tohuman influence. So, intense hunting pressuresover 10,000 years might directly influence local

extinctions of faunal populations and probably -still unknown- contributing to shape the structureof current primate populations as might betentatively inferred from the results. There iscurrent evidence to support this idea. For instance,Souza-Mazurek et al. (2000 579; 591) indicatedthat among the Waimiri-Atroari in northernBrazil, “sex ratios of spider monkeys [Atelespaniscus] killed were heavily biased towardsfemales indicating a stronger hunting pressure onthose individuals.” Thus, it seems that potentialsex-based differences in hunting selection areaffecting the group structure of feral Neotropicalprimates groups; naturally more detailed studiesto evaluate the effect of selective hunting on theprimate population dynamic are needed.Moreover, Peres (1991) described localextinctions of woolly monkey (Lagothrix sp.)populations as a consequence of intense huntingin the Amazonia. He said that woolly monkeypopulation density (number/km2) varied from 0and 7 in hunted sites to 17 and 30 in non-hunted sites. In addition, Creole populations inLatin America intensively hunt primates formarket networks that expand the limits of huntingfrom family consumption to hunting forcommercial purposes (Mittermeier, 1987, forunderstanding Amazonian Caboblos -BrazilianCreoles- economy: Nugent, 1993). Bush meat is apreferred food source in the local marts. Forexample, Castro et al. (1975) found that in sixmonths, in the popular market of Iquitos, Peru,were sold 1,700 Lagothrix sp., 1,396 Cebus sp.,557 Alouatta sp., 321 Pithecia sp. and 198 Atelessp. (approximate calculation by B. U.). Thus,together with the Congo Basin in Africa where thebush meat crisis is aggressively threatening primatepopulations (Peterson, 2003), Amazonia appearsto be the other geographical area critically threatenby hunting (Mittermeier, 1987; see Results).

Then, to the question, why New Worldprimates adopt the above indicated anti-predatory behaviors even if they are not efficientagainst their main predator, Homo sapiens?. Twodimensions might be playing a role on it. First,humans have interacted with New Worldmonkeys during just a short timeframe (~11,000years) compared to the long-time period sinceprimates had been found in the New World; thelate Oligocene when Branisella boliviana

B. Urbani 95

inhabited the central part of South America(Kay et al. 2002). On the other hand, humansocial practices and material culture (particularlyweapons) take part in the equation of predation ofNeotropical monkeys, being factors unique toHomo in the Neotropics. Nevertheless, extensivefield research is needed to properly address thisissue.

Probably one of the best ways to inquiryabout the relationship between Homo and NewWorld primates, including in particular hunting-predatory- practices, is to look into newinsights from ethnoprimatological perspectiveson the conceptualization of monkeys byAmerindians; a research line that have been justinitiated after Sponsel (1997) (in theNeotropics: Fleck et al., 1999; Cormier, 2002,2003; Lizarralde, 2002; Shepard, 2002; Urbani& Gil, 2001). Many Amerindian groups havespecial tied relations with monkeys, andassociated and classified them as “beings likehumans” (e.g. Kalapalo: Basso, 1973). Forinstance, among the Mekranoti, a Tupi languageAmerindian group related to the Kayapo group,indicated that other Amerindian groups used theword “Kaya-po” for referring them; the word“Kayapo” means the people that “resembledmonkeys” (Werner, 1985: 173). Ethnoprimatologicalstudies are fundamental for understanding therelationship between human and non-humanprimates. Moreover, these works will be essentialfor understanding the past and present inter-connection of both sympatric primates and evenmore, to comprehend the still unknown butcritical future of them. In addition, at themoment there are few systematic mentionsdescribing the New Word primate reactions tohuman primates, and in principle there arepractically no works specifically describing howhumans have potentially affected social andgrouping behaviors in Neotropical monkeys (forOld World primates: Coppinger & Maguire,1980; Bshary, 2001; Tenaza, 1990; Tenaza &Tilson, 1985). This is an interesting area for futureprimatological research in the tropical Americas.Moreover, recent data on naïve behaviors andgroup composition of wild primates (forchimpanzees: Morgan & Sanz, 2003) may befundamental for comparing with primatepopulations subjected to often human contact

(including primatologists), in order to understandpotential differences in primate social structuredue to human presence or impact.

Acknowledgments

Many thanks to Loretta A. Cormier, ManuelLizarralde, Leslie E. Sponsel and Robert S. Voss forsending me their publications for a previousbibliographic review (Urbani, 2002) that wereuseful for thinking in this work. To the UIUClibrary staff for make available valuable help inreferences searching. To Paul Garber and theanonymous referees for their critical suggestions. ToTania for her passion in the conservation of tropicalrainforests and her enormous love. B. Urbani isgranted by a Fulbright-OAS Scholarship.

References

Alvard M. 1993. Testing the “Ecologically noblesavage” hypothesis: Interpecific prey choiceby Piro hunters of Amazonian Perú. Hum.Ecol., 21: 355-387.

Alvard M. 1995. Intraspecific prey choice byAmazonian hunters. Curr. Anthropol., 36:789-818.

Alvard M. 1995. Shotguns and sustainablehunting in the neotropics. Oryx, 29: 58-66.

Alvard M. & Kaplan H. 1991. Procurementtechnology and prey mortality amongindigenous neotropical hunters. In M.C.Stiner (ed): Human predators and preymortality, pp. 79-104. Westview Press,Boulder.

Århem K. 1976. Fishing and hunting among theMakuma: Economy, ideology and ecologicaladaptation in the northwest Amazon. Ann.Report of the Gotenborgs Etnografiska Museum,: 27-44.

Baer P. & Merrifield W.R. 1972. Los Lacandones deMéxico. Dos Estudios. Instituto NacionalIndigenista, Secretaría de Educación Pública,Mexico City.

Baker M. 1992. Capuchin monkeys (Cebuscapucinus) and the Ancient Maya. AncientMesoam., 3: 219-228.

Baleé W. 1985. Ka’apor ritual hunting. Hum.


Ecol., 13: 485-510. Basso E.B. 1973. The Kalapalo Indians of Central

Brazil. Holt, Rinehart and Winston, NewYork.

Beckerman S. 1979. The abundance of protein inAmazonia: A reply to Gross. Am. Anthopol.,81: 53-56.

Beckerman S. 1980. Fishing and hunting by theBari in Colombia. Working Papers on SouthAmerican Indians, 2: 68-109.

Beckerman S. & Sussenbach T. 1983. Aquantitative assessment of the dietarycontribution of game species to thesubsistence of South American Tropical foresttribal peoples. In J. Clutton-Brock & C.Crigson (eds): Animals and Archaeology. 1.Hunters and their preys, pp. 337-350. BARInternational Series 163, Oxford.

Berlin B. & Berlin E.A. 1983. Adaptation andethnozoological classification: Theoreticalimplications of animal resources and diet ofthe Aguaruna and Huambisa. In R.B. Hames& W.T. Vickers (eds): Adaptive responses ofnative Amazonians, pp. 301-325. AcademicPress, New York.

Blomberg R. 1956. The naked Aucas. An account ofthe Indians of Ecuador.: George Allen &Unwin, London.

Boinski S. & Chapman C.A. 1995. Predation onprimates: Where are we and what’s next?.Evol. Anthropol., 4: 1-3.

Boinski S.; Treves A. & Chapman C.A. 2000. Acritical evaluation of the influence ofpredators on primates: Effect on group travel.In S. Boinski & P.A. Garber, (eds): On theMove. How and why animals travel in groups,pp. 43-72. University of Chicago Press,Chicago.

Braakhuis H.E.M. 1987. Artificers of the Days:Functions of the Howler Monkey Godsamong the Mayas. Bijdragen tot de Taal-,Land-, en Volkenkunde 143. FlorisPublications, Dortrechs.

Bshary R. 2001. Diana monkeys, Cercopithecusdiana, adjust their anti-predator responsebehaviour to human hunting strategies.Behav. Ecol. Sociobiol., 50: 251-256.

Bush M.B. & Colinvaux P.A. 1991. A pollenrecord of a complete glaciar cycle fromlowland Panama, J. Vegetation Sci., 1: 105-118.

Campos R. 1977. Producción de pesca y caza enuna comunidad Shipibo en el río Pisqui.Amaz. Peruana, 1: 53-74.

Carneiro R.L. 1970. Hunting and hunting magicamong the Amachuaca of the PeruvianMontaña. Ethnology, 9: 331-341.

Carpenter C.R. 1934. A field study of thebehaviour and social relations of howlingmonkeys. Comp. Psychol. Monogr., 10: 1-168.

Castro N., Revilla J. & Neville M. 1975. Carne demonte como una fuente de proteínas enIquitos, con referencia especial a monos. Rev.Forestal del Perú. 5: 19-32. Reprinted in: N.Castro-Rodríguez (ed): La primatología en elPerú, pp. 17-35. 1990. Proyecto Peruano dePrimatología, Lima.

Chapman C.A. 1986. Determinants of group sizein primates: The importance of travel cost. InS. Boinski & P.A. Garber (eds): On the Move.How and why animals travel in groups, pp. 23-42. University of Chicago Press Chicago.

Chapman C.A. & Peres C.A. 2001. Primateconservation in the New Millenium: The roleof scientists. Evol. Anthropol., 10: 16-33.

Chapman C. A. 2000. Boa constrictor predationand group response in white-faced Cebusmonkeys. Biotropica, 18: 171-172.

Cheney S. & Wragham R. 1987. Predation. InB.B. Smuts, R.M. Cheney, R.M. Seyfarth,R.W. Wrangham & T.T. Struhsaker (eds):Primate societies, pp. 227-239. University ofChicago Press, London.

Chinchilla F. A. 1997. La dieta del jaguar(Panthera onca), el puma (Felis concolor), y elmanigordo (Felis pardalis) (Carnivora:Felidae) en el Parque Nacional Corcovado,Costa Rica. Rev. Biol. Trop., 45: 1223-1229.

Coimbra-Fihlo A.F. & Mittermeier R.A. 1981.Ecology and Behavior of Neotropical Primates.Academia Brasileira de Ciências, Rio deJaneiro.

Colinvaux P.A. & Bush M.B. 1991. The rain-forest ecosystems as a resource for huntingand gathering. Am. Anthropol., 93: 153-160.

Conzemius E. 1984. Estudio etnográfico sobre losindios Miskitos y Sumus de Honduras yNicaragua. Libro Libre, San José.

Coppinger R.P. & Maguire J.P. 1980.Cercopithecus aethiops of St. Kitts: Apopulation estimate based on human

B. Urbani 97

predation. Caribb. J. Sci., 15: 1-7.Cormier L.A. 2002. Monkey as food, monkey as

child: Guajá symbolic cannibalism. In A.Fuentes & L.D. Wolfe (eds): Primates face toface: Conservation implications of human andnonhuman primate interconnections, pp. 63-84. Cambridge University Press, Cambridge.

Cormier L.A. 2003. Kinship with monkeys. TheGuajá foragers of Eastern Amazonia. ColumbiaUniversity Press, New York.

Côrrea H.K.M & Coutinho P.E.G. 1997. Fatalattack of a pit viper, Bothrops jararaca, on aninfant buffy-tufted ear marmoset (Callithrixaurita). Primates, 38: 215-217.

Cowlishaw G. & Dunbar R. 2000. PrimateConservation Biology. University of ChicagoPress, Chicago.

Crocket J.C. 1985. Vital souls. Bororo cosmology,natural symbolism, and shamanism. Universityof Arizona Press, Tucson.

Cuarón A.D. 1997. Conspecific aggression andpredation: Costs for a solitary mantled howlermonkey. Folia Primatol., 68: 100-105.

Descola, P. 1996. The spears of twilinght. Life anddeath in the Amazon jungle. The New Press,New York.

Di Fiore A. 2002. Predator sensitive foraging inateline primates. In L.E. Miller, (ed): Eat or beEaten, pp. 242-268. Cambridge UniversityPress, Cambridge.

Eakin E., Lauriault E. & Boonstra H. 1980.Bosquejo etnográfico de los Shipibo-Conibo delUcayali.: Ignacio Prado Pastor Ediciones,Lima.

Eason P. 1989. Harpy eagle attempts predation onadult howler monkey. Condor, 91: 469-470.

Emmons L.J. 1987. Comparative feeding ecologyof felids in a Neotropical rainforest. Behav.Ecol. Sociobiol., 20, 271-283.

Ferrari S.F., Pereira W., Santos R.R. & Viega L.M.2002. Ataque fatal de uma jibóia (Boa sp.) emum cuixú (Chiropotes satanas utahicki). Livrode Resumos do X Congresso Brasileiro dePrimatologia: Amazonia, a última fronteira, 66.

Fleck D.W., Voss R.S. & Patton J.L. 1999.Biological basis of saki (Pithecia) folk speciesrecognized by the Matses Indians ofAmazonian Peru. Int.. J. Primatol., 20: 1005-1028.

Fowler J.M. & Cope J.B. 1964. Notes on the harpy

eagle in British Guiana. The Auk, 81: 257-273.Fuentes A. & Wolfe L. D. (eds) 2002. Primates

face to face: Conservation implications ofhuman and nonhuman primateinterconnections. Cambridge University Press,Cambridge.

Galef B.G.Jr., Mittermeier R.A. & Bailey R.C.1976. Predation by the tayra (Eira barbara). J.Mammal., 57: 760-761.

Garber P.A. & Bicca-Marques J.C. 2002. Evidenceof predator sensitive foraging and traveling insingle- and mixed-species tamarin troops. InL.E. Miller (ed): Eat or be Eaten., pp. 138-153. Cambridge University Press, Cambridge.

García del Cueto H. 1989. Acerca de laconnotación simbólico-ritual del mono en lasociedad prehispánica (Altiplano Central). InA. Estrada, R. López-Wilchis & R. Coates-Estrada (eds): Primatología en México:Comportamiento, ecología, aprovechamiento yconservación de primates, pp. 144-159.Universidad Autónoma Metropolitana-Iztapalapa, Mexico City.

Gilbert K.A. 2000. Attempted predation on awhite-faced saki in the Central Amazon.Neotrop. Primates, 8: 103-104.

Goldizen A.W. 1987. Tamarins and marmosets:Communal care of offspring. In B.B. Smutts,R.M. Cheney, R.M. Seyfarth, R.W.Wrangham & T.T. Struhsaker (eds): Primatesocieties, pp. 34-43. Chicago University Press,London.

González-Kirchner J.P. & Sainz de la Maza M.1998. Primates hunting by GuaymiAmerindians in Costa Rica. Human Evol., 13:15-19.

Gregor T. 1977. Mehinaku. The drama of daily lifein a Brazilian Indian village.: University ofChicago Press, Chicago.

Gross D. 1975. Protein capture and culturaldevelopment in the Amazon basin. Am.Anthopol., 77: 526-549.

Hames R.B. 1979. A comparison of theefficiencies of the shotgun and the bow inneotropical forest hunting. Human Ecol., 7:219-252.

Harner M. J. 1972. The Jivaro: people of the SacredWaterfalls. Doubleday-Natural History, NewYork.

Heckenberger M.J., Kuikuro A., Kuikuro U.T.,


Russell J.C., Schmidt M., Fausto C. &Franchetto B. 2003. Amazonia 1492: PristineForest or Cultural Parkland?. Science, 301:1710-1714.

Heymann E.W. 1987. A field observation ofpredation on a moustached tamarin (Saguinusmystax) by an anaconda. Int. J. Primatol., 8:193-195.

Hill K. & Padwe J. 2000. Sustainability of Achehunting in the Mbaracayu Reserve, Paraguay.In J.G. Robinson & E.L. Bennett (eds):Hunting for sustainability in tropical forests, pp.79-105. Columbia University Press, New York.

Hill R.A. & Dunbar R.L.M. 1998. An evaluationof the roles of predation rate and predationrisk as selective pressures on primate groupingbehaviour. Behaviour, 135: 411-430.

Isbell L.A. 1994. Predation on primates:Ecological patterns and evolutionaryconsequences. Evol. Anthropol., 3: 61-71.

Izawa K. 1978. A field study of the ecology andbehavior of the black-mantled tamarin(Saguinus nigricollis). Primates, 19: 241-274.

Izor R.J. 1985. Sloths and other mammalians preyof the harpy eagle. In G. G. Montgomery,(ed): The evolution and ecology of armadillos,sloths and vermilinguas, pp. 343-346.Smithsonian Institution Press, Washington.

Janson C.H. 1998. Testing the predationhypothesis for vertebrate sociality: Prospectsand pitfalls. Behaviour, 135: 389-410.

Julliot C. 1994. Predation of a young spider monkey(Ateles paniscus) by a crested eagle (Morphnusguianensis). Folia Primatol., 63: 75-77.

Kappeler P.M. & Pereira M.E. 2003. Appendix. Aprimate life history database. In P.M.Kappeler & M.E. Pereira (eds): Primate lifehistories and socioecology, pp. 313-330.University of Chicago Press, Chicago.

Karadimas D. 1999. La constellation des quatresignes. Interprétation ethno-archéoastronomique des motifs de “ElCarchi-Capulí” (Colombie, Équator). J. Soc.Américanistes, 85: 115-145.

Kay R.F., Williams B.A. & Anaya F. 2002. Theadaptation of Branisella boliviana, the earliestSouth American monkey. In Plavcan J.M.,Kay R.F., Jungers W.L. & C.P. van Schaik,(eds): Reconstructing behavior in the primatefossil record, pp. 339-370. Kluwer

Academic/Plenum Publishers, New York.Kensinger K.M., Rabineau, P., Tanner, H.,

Ferguson, S.G. & Dawson A. 1975. TheCashinahua of Eastern Peru. Studies inAnthropology and Material Culture, Vol. 1.The Haffereffer Museum of Anthropology,Brown University Press, Providence.

Kinzey W.G. (ed) 1997. New World Primates:Ecology, evolution and behavior. Aldine deGruyter, New York.

Kracke W.H. 1978. Force and persuasion.Leadership in an Amazonian Society. Universityof Chicago Press, Chicago.

Lizarralde M. 2002. Ethnoecology of monkeysamong the Barí of Venezuela: perception, useand conservation. In A. Fuentes & L.D.Wolfe (eds): Primates face to face: Conservationimplications of human and nonhuman primateinterconnections, pp. 85-100. CambridgeUniversity Press, Cambridge.

Lizot J. 1979. On food taboos and Amazon culturalecology. Curr. Anthropol., 20: 150-151.

MacPhee, G. & I. Horovitz. 2002. ExtinctQuaternary platyrrhines of the greaterAntilles and Brazil. In W.C. Hartwig (ed):The primate fossil record, pp. 189-200.Cambridge University Press, Cambridge.

Maybury-Lewis D. 1967. Akwê-Shavante society.Clarendon Press, Oxford.

McDonald D.R. 1977. Food taboos: A primitiveenvironmental protection agency (SouthAmerica). Anthropos, 72: 735-748.

Meggers B.J. 1971. Amazonia. Man and culture ina counterfeit paradise. Aldine-Atherton, NewYork.

Mena V.P., Stallings J.R., Regalado B.J. & CuevaL.R. 2000. The sustainability of currenthunting practices by the Huaorani. In J.G.Robinson & E.L. Bennett (ed): Hunting forsustainability in tropical forests, pp. 57-78.Columbia University Press, New York.

Merriam J.C. 1998 - Community wildlifemanagement by Mayangna Indians in theBosawas Reserve, Nicaragua. M. S. thesis,Idaho State University.

Miller L.E. 2002. An introduction to predatorsensitive foraging. In L. E. Miller (ed): Eat orbe Eaten, pp. 1-20. Cambridge UniversityPress, Cambridge.

Milton K. 1991. Comparative aspects of diet in

99B. Urbani

Amazonian forest-dwellers. In: A. Whiten &E.M. Widdowson (eds): Foraging strategies andnatural diet of monkeys, apes and humans, pp.93-103. Clarendon Press, Oxford.

Mitchell C.L., Boinski, S. & van Schaik, C.P.1991. Competitive regimes and femalebonding in two species of squirrel monkey(Saimiri oerstedi and S. sciureus). Behav. Ecol.Sociobiol., 28: 55-60.

Mittermeier R.A. 1987. Effects of hunting on rainforest primates. In C.W. Marsh & R.A.Mittermeier (ed): Primate conservation in thetropical rain forest, pp. 109-146. Alan R. Liss,New York.

Mittermeier R.A. 1991. Hunting and its effect onwild primates populations in Suriname. InJ.G. Robinson & K.H. Redford (eds):Neotropical wildlife use and conservation, pp.93-107. The University of Chicago Press,Chicago.

Mittermeier R.A., Rylands A.B., Coimbra-Fihlo,A.F. & da Fonseca, G.A.B. 1988. Ecology andbehavior of Neotropical Primates. WorldWildlife Fund, Washington.

Montgomery E.I. 1970. With the Shiriana inBrazil.: Kendall/Hunt Publishing, Dubuque.

Morgan D. & Sanz C. 2003. Naïve encounterswith chimpanzees in the Goualougo Triangle,Republic of Congo. Int. J. Primatol., 24: 369-381.

Moynihan M. 1970. Some behavior patterns ofPlatyrrhine monkeys, II. Saguinus geoffroyiand some other tamarins. SmithsonianContrib. Zool., 29: 1-77.

Murphy R.F. & Quain B. 1966. The TrumaíIndians of Central Brazil. Monographs of theAmerican Ethnological Society, 24. Universityof Washington Press, Seattle.

Nimuendajú C. 1946. The Eastern Timbira.University of California Press, Berkeley.

Nugent S. 1993. Amazonian Caboclo society: Anessay on invisibility and peasant economy. BergPublishers, Oxford.

Olmos F. 1994. Jaguar predation on muriqui,Brachyteles arachnoids. Neotrop. Primates, 2: 16.

Oversluijs-Vasquez M.R. & Heymann E.W. 2001.Crested eagle (Morphnus guianensis)predation on infant tamarins (Saguinusmystax and Saguinus fuscicollis,Callitrichinae). Folia Primatol., 72: 301-303.

Peres C.A. 1990. A harpy eagle successfullycaptures an adult male red howler monkey.Wilson Bull., 102: 560-561.

Peres C.A. 1991. Humboldt’s woolly monkeysdecimated by hunting in Amazonia. Oryx 25:89-95.

Peterson, D. 2003. Eating Apes. University ofCalifornia Press, Berkeley.

Redford K.H. 1992. The empty forests.BioScience, 42: 412-422.

Reichel-Dolmatoff G. 1989. Amazonian Cosmos.The sexual and religious symbolism of theTukano Indians. University of Chicago Press,Chicago.

Rettig N.L. 1978. Breeding behavior of the harpyeagle, Harpia harpyja. The Auk, 95: 629-643.

Reverte J.M. 1967. Los indios Teribes de Panamá.Talleres de la Estrella de Panamá, PanamaCity.

Robinson J.G. 1993. The limits to caring:Sustainable living and the loss of biodiversity.Conserv. Biol., 7: 20-28.

Robinson J.G. & Redford K.H. 1994. Measuringthe sustainability of hunting in tropicalforests. Oryx, 28: 249-256.

Roosevelt A.C., Lima da Costa M., LopesMachado C., Michab M., Mercier N.,Valladas H., Feathers J., Barnett W., Imazioda Silveira M., Henderson A., Sliva J.,Chernoff B., Reese D.S., Holman J.A., TothN. & Schick K. 1996. Paleoindian cavedwellers in the Amazon: The peopling of theAmericas. Science, 272: 373-384.

Ross E.B. 1978. Food taboos, diet, and huntingstrategy: The adaptation to animals inAmazon cultural ecology. Curr. Anthropol.,19: 1-36.

Saffirio G. & Scaglion R. 1982. Huntingefficiency in acculturated Yanomama villages.J. Anthropol. Res., 38: 315-328.

Schaller G.B. 1983. Mammals and their biomasson a Brazilian Ranch. Arq. Zool. (São Paulo),31: 1-36.

Seeger A. 1981. Nature and society in CentralBrazil. The Suya Indians of Mato Grosso.Harvard University Press, Cambridge.

Setz E.Z.F. 1991. Animals in the Nambiquaradiet: methods of collection and processing. J.Ethnobiol., 11: 1-22.

Shepard G.Jr. 2002. Primates in Matsigenka


subsistence and worldview. In A. Fuentes &L. D. Wolfe (eds): Primates face to face:Conservation implications of human andnonhuman primate interconnections, pp. 101-136. Cambridge University Press,Cambridge.

Sherman P.T. 1991. Harpy eagle predation on a redhowler monkey. Folia Primatol., 56: 53-56.

Simpson, J.A. & Weiner E.S.C. 1989. The OxfordEnglish Dictionary. Volume VII. ClarendonPress, Oxford.

Siskind J. 1973. To hunt in the morning. OxfordUniversity Press, New York.

Souza-Mazurek R., Pedrinho T., Feliciano X.,Hilario W., Geroncio S. & Marcelo E. 2000.Subsistence hunting among the WaimiriAtroari Indians in central Amazonia, Brazil.Biodivers. Conserv., 9: 579-596.

Sponsel L.E. 1997. The human niche inAmazonia: Explorations in ethnoprimatology.In W.G. Kinzey (ed): New World primates.Ecology, evolution, and behavior, pp. 143-165.Aldine de Gruyter, New York.

Stanford, G.B. 1998. Chimpanzee and red colobus.The ecology of predator and prey. HarvardUniversity Press, Cambridge.

Stafford B.J. & Murad-Ferreira, F. 1995.Predation attempts on Callitrichids in theAtlantic coastal rain forest of Brazil. FoliaPrimatol., 65: 229-233

Struhsaker T.T. 1975. The red colobus monkey.Chicago University Press, Chicago.

Tenaza R.R. 1990. Effects of human predation onbehavior of pig-tailed langurs in the MentawaiIslands. Am. J. Primatol., 20: 237-238.

Tenaza R.R, Tilson R.L. 1985. Human predationand Kloss’s gibbon (Hylobates klossii) sleepingtrees in Siberut Island, Indonesia. Am. J.Primatol., 8: 299-308.

Terborgh J. 1983. Five New World Primates.Princeton University Press, Princeton.

Thompson, J.E. 1930. Ethnology of the Mayas ofsouthern and central British Honduras. FieldMuseum of Natural History, Publication 274.Anthropological series, 27: 27-213.

Townsend W.R. 2000. The sustainability ofsubsistence hunting by the Siriono Indians of

Bolivia. In J.G. Robinson & E.L. Bennett(eds): Hunting for sustainability in tropicalforests pp. 267-281. Columbia UniversityPress, New York.

Tsukahara T. & Nishida T. 1990. The firstevidence for predation by lions on wildchimpanzees. XIIIth Congress of theInternational Primatological Society Abstracts(Nagoya), pp. 82

Urbani B. 2002. Neotropical ethnoprimatology:An annotated bibliography. Neotrop.Primates, 10: 24-26.

Urbani B. & Gil E. 2001. Consideraciones sobrerestos de primates de un yacimientoarqueológico del Oriente de Venezuela(América del Sur), Cueva del Guácharo,estado Monagas. Munibe, 53: 135-142.

van Akkeren R. 1998. The monkey and the blackheart. Polar North in Ancient Mesoamerica.In A.L. Izquierdo (ed): Memorias del TercerCongreso Internacional de Mayistas, pp. 165-185. Universidad Nacional Autónoma deMéxico, Mexico City.

van Schaik C.P., Horstermann M. 1994. Predationrisk and the number of adult males in aprimate group: A comparative test. Behav.Ecol. Sociobiol., 35: 261-272.

Vickers W. 1980. An analysis of Amazonian huntingyields as a function of settlement age. WorkingPapers on South American Indians, 2: 7-30.

Viveiros de Castro E. 1992. From the enemies pointof view. Humanity and divinity in anAmazonian society. University of ChicagoPress, Chicago.

von Graeve B. 1989. The Pacaa Nova. Clash ofcultures on the Brazilian border. BroadviewPress, Petesburgh.

Wagley C. 1977. Welcome of tears. The TapirapéIndians of Central Brazil. Oxford UniversityPress, New York.

Wagley C. &, Galvão E. 1949. The TeneteharaIndians of Brazil. A culture in transition.Columbia University Press, New York.

Werner D. 1985. Amazon Journey. Ananthropologist’s year among Brazil’s MekranotiIndians. Simon and Schuster, New York.

101B. Urbani


A

pp

en

dix

- C

om

pari

son

of

pre

dati

on

on

New

Wo

rld

pri

mate

s.

B. Urbani 103

Ap

pen

dix

- (

con

tin

ued

).


Ap

pen

dix

- (

con

tin

ued

).

B. Urbani 105

Ap

pen

dix

- (

con

tin

ued

).

106 Predation on Neotropical MonkeysA

pp

en

dix

- (

con

tin

ued

).

B. Urbani 107

Ap

pen

dix

- (

con

tin

ued

).


Ap

pen

dix

- (

con

tin

ued

).

B. Urbani 109

Ap

pen

dix

- (

con

tin

ued

).

the targeted monkey: a re-evaluation of predation on new world primates vol83/urbani.pdf ·...

Documents