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

JASs Reports
Journal <strong>of</strong> Anthropological Sciences
Vol. 83 (2005), pp. 89-109
<strong>The</strong> <strong>targeted</strong> <strong>monkey</strong>: a <strong>re</strong>-<strong>evaluation</strong> <strong>of</strong> p<strong>re</strong>dation on
New World primates
Bernardo Urbani
Department <strong>of</strong> Anthropology, University <strong>of</strong> Illinois at Urbana-Champaign, 109 Davenport Hall, 607 S
Mathews Ave., Urbana, Illinois 61801, USA, e-mail: burbani@uiuc.edu
Summary – This work <strong>re</strong>views the information <strong>re</strong>lated to p<strong>re</strong>dation on Neotropical primates by human
and non-human p<strong>re</strong>dators. Paradoxically, humans have been systematically neglected while evaluating the
potential effects <strong>of</strong> p<strong>re</strong>dation in the structu<strong>re</strong> <strong>of</strong> non-human primate populations. P<strong>re</strong>dation paradigms do not
include humans in their propositions. In this <strong>re</strong>view, it is shown that effectively humans a<strong>re</strong> the main p<strong>re</strong>dators
<strong>of</strong> <strong>monkey</strong> communities in the Neotropics. <strong>The</strong> <strong>re</strong>sults also suggest that humans do not fit with p<strong>re</strong>dation
theo<strong>re</strong>tical views given for non-human p<strong>re</strong>dators. Homo cultural hunting practices contribute to this
situation. For example, humans seem to p<strong>re</strong>fer larger primate groups that allow their location for hunting, or
humans tend to p<strong>re</strong>y primary larger <strong>monkey</strong>s with longer interbirth interval. In sum, it is suggested that since
at least 11,000 years <strong>of</strong> human occupation in the Neotropics, Homo might have been played a fundamental
role in the cur<strong>re</strong>nt organization and distribution <strong>of</strong> primate populations, including local extinctions. Humans
seemed to have potentially influenced New World primate population in such short ecological time scale.
Keywords – P<strong>re</strong>datory behavior, ethnoprimatology, human hunting, p<strong>re</strong>dation paradigms,
transdisciplinary perspective, conservation, Latin America.
Introduction
“<strong>The</strong> indications a<strong>re</strong> that man is the most
serious enemy <strong>of</strong> howlers and that occasionally
young animals may be attacked by ocelots” said
Cla<strong>re</strong>nce Ray Carpenter in 1934 after observing
mantled howler <strong>monkey</strong>s in Panama (Carpenter,
1934: 129). In fact, it was his last conclusion in
the first systematic primate behavioral <strong>re</strong>search
conducted in the wild, and probably also the first
scientific account <strong>of</strong> the impact <strong>of</strong> p<strong>re</strong>dators on
feral primates. After this work, other <strong>re</strong>searches
have taken p<strong>re</strong>dation risk into account as a
potential factor influencing the evolution <strong>of</strong>
sociality in general, and the social structu<strong>re</strong> <strong>of</strong>
primate populations (e.g. Cheney & Wragham,
1987). For example, since the 1960s chimpanzees
had been the subjects <strong>of</strong> the longest-term studies
even carried out for any wild mammal; but
paradoxically it was not until 1990, that the first
case <strong>of</strong> p<strong>re</strong>dation on chimpanzees by lions was
<strong>re</strong>ported (Tsukahara & Nishida 1990). In
addition, p<strong>re</strong>dation has been systematically cited in
works <strong>of</strong> the natural history <strong>of</strong> primates (e. g.
Kinzey, 1997); however, <strong>re</strong>ports <strong>of</strong> p<strong>re</strong>dation a<strong>re</strong>
ext<strong>re</strong>me scarce. In part, this <strong>re</strong>lates to difficulties
obtaining field data on p<strong>re</strong>dation because <strong>of</strong> its
low rate and rapid occur<strong>re</strong>nce. In addition, Isbell
(1994) suggested that limited p<strong>re</strong>dation
observation might be <strong>re</strong>lated to the observers (field
primatologists) that a<strong>re</strong> normally not p<strong>re</strong>sent
when main p<strong>re</strong>dators a<strong>re</strong> active (at night) and the
p<strong>re</strong>sence <strong>of</strong> the observer may inhibit the
appearance <strong>of</strong> p<strong>re</strong>dators during the day.
However, p<strong>re</strong>dation and p<strong>re</strong>dation risk have
been conside<strong>re</strong>d important in modeling the
structu<strong>re</strong> and organization <strong>of</strong> the extant and
extinct primate populations (see <strong>re</strong>view: Isbell,
1994). In this sense, it has been argued that some
“anti-p<strong>re</strong>datory” behaviors such as vigilance,
polyspecific associations, group cohesion, di<strong>re</strong>ct or
active defense and alarm calls a<strong>re</strong> <strong>re</strong>lated to the
selective p<strong>re</strong>ssu<strong>re</strong> <strong>of</strong> p<strong>re</strong>dation on primates
(Cheney & Wragham, 1987; Isbell, 1994). Van
Schaik & Horstermann (1994) proposed a
hypothesis that suggested that the quantity <strong>of</strong>
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90 P<strong>re</strong>dation on Neotropical Monkeys
primate males -and group composition- is not
only <strong>re</strong>lated to sexual competition but also to
p<strong>re</strong>dation risk. <strong>The</strong>ir hypothesis p<strong>re</strong>dicts that
males a<strong>re</strong> mo<strong>re</strong> vigilant than females, and play a
g<strong>re</strong>ater role in p<strong>re</strong>dator detection.
In her literatu<strong>re</strong> <strong>re</strong>view, Isbell (1994: 65-68)
characterized five patterns <strong>of</strong> p<strong>re</strong>dation on
primates. <strong>The</strong>se patterns a<strong>re</strong>, 1) larger primates a<strong>re</strong>
expected to be less vulnerable to p<strong>re</strong>dation than
a<strong>re</strong> smaller primates; 2) an individual’s
vulnerability to p<strong>re</strong>dation inc<strong>re</strong>ased in unknown
or unfamiliar a<strong>re</strong>as (for a definition <strong>of</strong>
vulnerability, see below: Miller, 2002); 3) primates
a<strong>re</strong> mo<strong>re</strong> vulnerable in the upper canopy, in
discontinuous fo<strong>re</strong>sts and on the fo<strong>re</strong>st edges than
in continuous, undisturbed fo<strong>re</strong>sts; 4) p<strong>re</strong>dation is
episodic and may be <strong>re</strong>lated to p<strong>re</strong>y p<strong>re</strong>fe<strong>re</strong>nces
and diffe<strong>re</strong>nt use <strong>of</strong> daily ranges and home ranges
between p<strong>re</strong>dators and p<strong>re</strong>y; and 5) ter<strong>re</strong>strial
primates have higher rates and risk <strong>of</strong> p<strong>re</strong>dation
than do arbo<strong>re</strong>al primates. In addition, Chapman
(2000) <strong>re</strong>viewed the p<strong>re</strong>dation avoidance
hypothesis by examining patterns <strong>of</strong> group
structu<strong>re</strong> and movement in 54 primate species. He
argued that living in groups, “[may] inc<strong>re</strong>ased
probability <strong>of</strong> p<strong>re</strong>dator detection, … [c<strong>re</strong>ate]
g<strong>re</strong>ater confusion <strong>of</strong> a p<strong>re</strong>dator trying to focus on
an individual p<strong>re</strong>y, … [propitiate] a dec<strong>re</strong>ased
probability <strong>of</strong> each individual being captu<strong>re</strong>d by
p<strong>re</strong>dators, … and inc<strong>re</strong>ased defense against
p<strong>re</strong>dators” (Chapman 2000: 27).
However, Janson (1998) indicated that the
<strong>evaluation</strong> <strong>of</strong> p<strong>re</strong>dation should be done with
caution due to the fact that p<strong>re</strong>sumed antip<strong>re</strong>datory
behaviors (e. g. alarm calls, vigilance,
living in larger groups) might be linked with other
behaviors. For instance, he indicated that for some
primate species it might be beneficial to live in
small groups that a<strong>re</strong> difficult for p<strong>re</strong>dators to
detect. In addition, Hill & Dunbar (1998)
suggested that p<strong>re</strong>dation risk is a mo<strong>re</strong>
comp<strong>re</strong>hensive parameter for evaluating p<strong>re</strong>dation
on primates than p<strong>re</strong>dation rate. <strong>The</strong>y argued that
potentially the role <strong>of</strong> p<strong>re</strong>dation as selective
p<strong>re</strong>ssu<strong>re</strong> might be mo<strong>re</strong> important for primates
when the risk <strong>of</strong> p<strong>re</strong>dation is perceived. In
addition, they indicated that p<strong>re</strong>dator-p<strong>re</strong>y
<strong>re</strong>lationship should be evaluated as the potentiality
that a given primate species might be <strong>re</strong>cove<strong>re</strong>d
from p<strong>re</strong>datory events by adjusting its behavior.
Recently, Miller (2002) added to the discussion
the concept <strong>of</strong> p<strong>re</strong>dation vulnerability in
understanding primate social organization. She
defined vulnerability as the “qualitative measu<strong>re</strong> <strong>of</strong>
the probability that an individual will be the victim
<strong>of</strong> a p<strong>re</strong>dator at any given moment” (Miller, 2002:
2). She indicated that this fact may play a major role
in the foraging strategies and decision-making
among primates. Miller (2002) suggested that
p<strong>re</strong>dation vulnerability might be evaluated using
th<strong>re</strong>e diffe<strong>re</strong>nt variables. A biological variable, in
which diffe<strong>re</strong>nt characteristics that a<strong>re</strong> “under
genetic control” and exp<strong>re</strong>ssed in the primate
phenotype might be significant for a given antip<strong>re</strong>datory
<strong>re</strong>sponse (e. g. body size); a social variable
such as diffe<strong>re</strong>nces in rank, group size and group
composition; and an environmental variable such
as the deg<strong>re</strong>e <strong>of</strong> p<strong>re</strong>dation vulnerability associated
with foraging location and the quality <strong>of</strong> the cover
or <strong>re</strong>fuge available to primates.
Humans as primate p<strong>re</strong>dators
Isbell (1994) and Cheney & Wrangham
(1987) briefly suggested that humans a<strong>re</strong> the
major p<strong>re</strong>dators on primates. Similarly, Boinski et
al. (2000: 47) stated in their <strong>re</strong>view, “primate
p<strong>re</strong>dators a<strong>re</strong> <strong>re</strong>stricted to nonhuman p<strong>re</strong>dators
because humans probably hunt most primates”
(italics a<strong>re</strong> mine). Thus, although the impact <strong>of</strong>
humans as p<strong>re</strong>dators on primates has been noted,
few <strong>re</strong>searchers have focused on the role <strong>of</strong> human
p<strong>re</strong>dators may have played in non-human primate
social organization (see: Sponsel, 1997). P<strong>re</strong>dation
by humans has been under-emphasized in the
theo<strong>re</strong>tical discussions about p<strong>re</strong>dation on nonhuman
primates. However, to hunt defined by the
Oxford English Dictionary (Simpson & Weiner,
1989: 496) is “the act <strong>of</strong> chasing wild animals for
the purpose <strong>of</strong> catching or killing them”; not less
than another form <strong>of</strong> p<strong>re</strong>dation. Consequently, as
indicated by Mittermeier (1987) and Chapman &
Pe<strong>re</strong>s (2001), New World primates a<strong>re</strong> highly
th<strong>re</strong>aten due to intense hunting or human
p<strong>re</strong>dation. Nevertheless, humans had <strong>of</strong>ten been
neglected in the literatu<strong>re</strong> on p<strong>re</strong>dation in
primates (Sponsel 1997). In this paper I will
examine primate p<strong>re</strong>y-human p<strong>re</strong>dator
<strong>re</strong>lationship in the Neotropics. So, I add<strong>re</strong>ss the
following questions,
a) Is the<strong>re</strong> evidence that p<strong>re</strong>dators affect the

structu<strong>re</strong> <strong>of</strong> primate communities in Neotropical
fo<strong>re</strong>sts?
b) How human p<strong>re</strong>dators differ from nonhuman
p<strong>re</strong>dators in p<strong>re</strong>y choice, hunting
techniques and potential affect on primate social
organization?
b) A<strong>re</strong> humans primary agents that influence
ecological changes such as local extinctions <strong>of</strong>
primate populations?
Methods
Information <strong>of</strong> p<strong>re</strong>dation on Neotropical
primates was compiled (see Appendix). Only
observed p<strong>re</strong>dation cases on <strong>monkey</strong>s by di<strong>re</strong>ct
sightings or from inspection <strong>of</strong> alimentary samples
such as primate skeletal material in p<strong>re</strong>dator nests
and primate <strong>re</strong>mains in p<strong>re</strong>dator feces we<strong>re</strong>
included. For this purpose, a <strong>re</strong>view <strong>of</strong> papers
<strong>re</strong>lated to p<strong>re</strong>dation and hunting on primates in
the Neotropics was systematically p<strong>re</strong>pa<strong>re</strong>d from
th<strong>re</strong>e bioanthropological -primatological- and
ethnographical sources.
First, a search was conducted <strong>of</strong> the major
anthropological and biological databases such as
Academic Search Elite, Anthropological Index,
Anthropological Literatu<strong>re</strong>, Biological Abstracts
(BIOSIS), Ecology Abstracts, JSTOR (including
35 ecological and anthropological journals dated
from 1867 to 1999), PrimateLit and Zoological
Record.
Secondly, a bibliographic compilation was
p<strong>re</strong>pa<strong>re</strong>d from the <strong>re</strong>views published by
McDonald (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 Fio<strong>re</strong> (2002), Fuentes & Wolfe (2002),
Urbani (2002) and Cormier (2003). Third, the
four main primatological journals American
Journal <strong>of</strong> Primatology (1981-2003), Folia
Primatologica (1963-2003), International Journal
<strong>of</strong> Primatology (1980-2003), Primates (1957-
2003), and the ethnological Journal <strong>of</strong>
Ethnobiology (1981-2003) we<strong>re</strong> also examined.
From the compiled <strong>re</strong>fe<strong>re</strong>nces, the following
information was obtained (see the Appendix):
a) Non-Homo p<strong>re</strong>dator: cor<strong>re</strong>sponds with non-
B. Urbani 91
human vertebrates that p<strong>re</strong>y on Neotropical
primates.
b) Homo p<strong>re</strong>dator: <strong>re</strong>fers only to Amerindian
hunters <strong>of</strong> <strong>monkey</strong>s in the tropical fo<strong>re</strong>sts. In this
work only Amerindians groups we<strong>re</strong> included,
considering the assumption that a<strong>re</strong> the human
populations that p<strong>re</strong>yed on <strong>monkey</strong>s inheriting
long-term traditional hunting techniques and
knowledge in the Neotropics (for Campesinos -
Latin American c<strong>re</strong>oles-, see discussion).
c) P<strong>re</strong>y: the <strong>targeted</strong> New World <strong>monkey</strong> genus
obtained by non-Homo or Homo p<strong>re</strong>dators.
Associated with this entry, in the cells <strong>of</strong> the
Appendix, information on the numbers <strong>of</strong>
observed p<strong>re</strong>dation cases, age/sex <strong>of</strong> the p<strong>re</strong>y, and
ranking <strong>of</strong> the p<strong>re</strong>y is collected. In this sense, 1)
Number <strong>of</strong> observed p<strong>re</strong>dation cases: <strong>re</strong>fer to the
quantity <strong>of</strong> the proved cases <strong>of</strong> p<strong>re</strong>dation on
<strong>monkey</strong>s; 2) Age/Sex: age and sex <strong>of</strong> the p<strong>re</strong>y; and
3) Ranking p<strong>re</strong>y: only for the cases <strong>of</strong> Homo
p<strong>re</strong>dators, <strong>re</strong>fers to the rank for a given primate
genus compa<strong>re</strong>d to all mammalian p<strong>re</strong>ys.
d) Total <strong>of</strong> p<strong>re</strong>y: the sum <strong>of</strong> all p<strong>re</strong>ys for a given
p<strong>re</strong>dator and the number <strong>of</strong> p<strong>re</strong>y in general.
e) Observation period: the total amount <strong>of</strong> time <strong>of</strong>
the primatological study or the time that the
ethnographer spent in the Amerindian
community in which the primate hunting events
we<strong>re</strong> observed.
f) F<strong>re</strong>quency <strong>of</strong> p<strong>re</strong>dation: <strong>re</strong>lationship between
the “Number <strong>of</strong> observed p<strong>re</strong>dation cases” per
hour. For both, primatological and ethnographical
data, in which the observation period was not
<strong>re</strong>ported by hour, I used the following conversion:
1 day = 10 hours, 1 month = 15 days, 1 year = 12
months. If the observation period was not
explicitly <strong>re</strong>ported, it is included as unknown.
g) Locality: site in which the observation was
done, including the <strong>re</strong>gion and the country.
Results
In Appendix, I summarizes the data archived
from the bibliographical search on hunting and
p<strong>re</strong>dation on Neotropical <strong>monkey</strong>s. A total <strong>of</strong> 89
entries we<strong>re</strong> obtained, 33 (37.1% for non-Homo
p<strong>re</strong>dators and 56 (62.9%) for Homo p<strong>re</strong>dators. For
non-Homo p<strong>re</strong>dators, 51.5% the events we<strong>re</strong>
<strong>re</strong>corded in non-Amazonian fo<strong>re</strong>sts (17/33, incl.
the Guianas, Central American and subtropic

92 P<strong>re</strong>dation on Neotropical Monkeys
fo<strong>re</strong>sts), and the other 48.5% (16/33) in the
Amazonia. In the case <strong>of</strong> Homo p<strong>re</strong>dators, 69.6%
(39/56) <strong>of</strong> the cases we<strong>re</strong> in Amazonia while
30.4% (17/56) we<strong>re</strong> in non-Amazonian fo<strong>re</strong>sts.
<strong>The</strong> f<strong>re</strong>quency <strong>of</strong> p<strong>re</strong>dation was quite similar from
0.045/hour for non-Homo p<strong>re</strong>dators to
0.048/hour in Homo p<strong>re</strong>dators (Appendix).
Among all non-Homo p<strong>re</strong>dators, avian
p<strong>re</strong>dators appear to be the most common. Of 32
cases, harpy eagles (Harpia harpija, 21.2%, 7/33)
and c<strong>re</strong>sted eagle (Morphnus guianensis, 12.1%,
4/33) a<strong>re</strong> the most common p<strong>re</strong>dators. Among
ter<strong>re</strong>strial non-Homo p<strong>re</strong>dators, jaguars (Panthera
onca) accounted for 15.2% (5/33) <strong>of</strong> the
observations. Ten other non-Homo p<strong>re</strong>dators
accounted for 51.5% (17/33) <strong>of</strong> p<strong>re</strong>datory events.
For non-Homo p<strong>re</strong>dators it was not possible
determine the sex/age classes <strong>of</strong> the <strong>monkey</strong>s
p<strong>re</strong>yed upon (69%, 29/42). However, for those
p<strong>re</strong>dation cases in which the information was
available, 21.4% (9/42) <strong>re</strong>p<strong>re</strong>sent young animals
(pooling together infants, juveniles and subadults)
and the <strong>re</strong>maining 9.5% (4/42) we<strong>re</strong> adults, while
69.1% we<strong>re</strong> <strong>of</strong> unknown age. In the few cases in
which the sex was observed, 9.5% (4/42) we<strong>re</strong>
males whe<strong>re</strong>as 7.1% (3/42) females, and the <strong>re</strong>st
83.4% <strong>re</strong>mained <strong>of</strong> unknown sex. For Homo
p<strong>re</strong>dators, in all cases the specific sex/age classes
we<strong>re</strong> unknown. In addition, as indicated in the
discussion (see below) 53.6% (30/56) <strong>of</strong> the
entries <strong>of</strong> Homo p<strong>re</strong>dators explicitly stated that
these human groups hunt at least one <strong>of</strong> the non-
Homo p<strong>re</strong>dators listed in Appendix. Mo<strong>re</strong>over, for
Homo p<strong>re</strong>dators it is important to identify the
ranking <strong>of</strong> <strong>monkey</strong> p<strong>re</strong>fe<strong>re</strong>nces among all
mammal p<strong>re</strong>y. It is inte<strong>re</strong>sting to note that among
Appendix entries, <strong>monkey</strong>s a<strong>re</strong> among the five
most common p<strong>re</strong>ys (27.8%, 42/151; see Tab. 1),
the <strong>re</strong>st 72.5% a<strong>re</strong> for primates p<strong>re</strong>fer<strong>re</strong>d over the
six position <strong>of</strong> p<strong>re</strong>fe<strong>re</strong>nce (15.9%, 24/151) or it is
unknown the choice rank among Amerindian
groups (56.3%, 85/151).
I tested the p<strong>re</strong>diction that Homo p<strong>re</strong>dators
have higher pr<strong>of</strong>iciency obtaining New World
<strong>monkey</strong>s than non-Homo p<strong>re</strong>dators. Pr<strong>of</strong>iciency
was defined as the number <strong>of</strong> successful hunts <strong>of</strong> a
given primate p<strong>re</strong>y. Both, Homo p<strong>re</strong>dators and
non-Homo p<strong>re</strong>dators we<strong>re</strong> compa<strong>re</strong>d using G-test.
<strong>The</strong> <strong>re</strong>sults indicate that humans a<strong>re</strong> mo<strong>re</strong>
pr<strong>of</strong>icient p<strong>re</strong>dators <strong>of</strong> New World primates (df=
12, p

population, it was suggested that a daily diet
should have at least 40-50 g/day per capita <strong>of</strong>
proteins whe<strong>re</strong> game meat and fish occupied an
important place apart <strong>of</strong> plant proteins from
sources such as mandioca (Gross, 1975, for
discussion see: Beckerman, 1979). For obtaining
Neotropical animal game, hunting practices a<strong>re</strong>
culturally based (Beckerman & Sussenbach, 1983;
Sponsel, 1997; Lizot, 1979; Lizarralde, 2002). For
example, Lizot (1979) indicated that food
p<strong>re</strong>fe<strong>re</strong>nces and taboos among Yanomami groups
determine the selection and consumption <strong>of</strong>
particular food items, particularly key game
animals. Also, cultural practices might influence
the inhibition for eating some <strong>monkey</strong>s. Among
the Desana <strong>of</strong> Amazonian Colombia, <strong>monkey</strong>s a<strong>re</strong>
prohibited meat for child<strong>re</strong>n, while howler
<strong>monkey</strong>s a<strong>re</strong> conside<strong>re</strong>d evil omens (Reichel-
Dolmat<strong>of</strong>f, 1971). In addition, humans a<strong>re</strong>
dramatically dec<strong>re</strong>asing the non-human p<strong>re</strong>dator
populations. For instance, over-hunting in the
Neotropics has c<strong>re</strong>ated the so-called “empty
fo<strong>re</strong>sts,” we<strong>re</strong> major mammals we<strong>re</strong> <strong>re</strong>moved from
their natural habitats (Redford, 1992; Robinson,
1993). <strong>The</strong> <strong>re</strong>sult is that in some tropical a<strong>re</strong>as,
humans maybe a<strong>re</strong> the only th<strong>re</strong>at to feral <strong>monkey</strong>
populations (Cowlishaw & Dunbar, 2000). This
has been documented in terms <strong>of</strong> the dec<strong>re</strong>asing
B. Urbani 93
woolly <strong>monkey</strong> population in the Brazilian
Amazon (Pe<strong>re</strong>s, 1991; see below).
On the other hand, humans do not fit well in
other proposed p<strong>re</strong>datory p<strong>re</strong>dictions <strong>of</strong> p<strong>re</strong>datorprimate
p<strong>re</strong>y interactions. In this sense, various
anti-p<strong>re</strong>datory behaviors such as mobbing, active
defense (e. g. throwing branches) and alarm calls
a<strong>re</strong> not successful for the <strong>monkey</strong>s because they
would be easily detected and killed by humans.
For example, some Amerindian groups such as the
Shirián <strong>of</strong> the upper Paragua River, Venezuela,
performed <strong>monkey</strong> calls in order to wait for an
acoustical <strong>re</strong>sponse to locate their potential p<strong>re</strong>y
(Urbani, pers. obs.). On the other hand, in
Mexico, the Lacandón people hear the roaring <strong>of</strong>
the <strong>monkey</strong>s from a long distance to chase and
hunt them (Baer & Merrifield, 1972). Mo<strong>re</strong>over,
other so-called anti-p<strong>re</strong>datory behaviors like
polyspecific associations and intragroup distance
might aid human hunters in locating the primate
groups. For instance, Balée (1985) <strong>re</strong>ported that
hunting howler <strong>monkey</strong>s by the Ka’apor in Brazil,
take them an average <strong>of</strong> just 50 minutes from the
moment they left from and <strong>re</strong>turned to the
settlement after the hunting party. This is the
smallest amount <strong>of</strong> time used for hunting any
mammalian game for this Amerindian group, the
next nea<strong>re</strong>st a<strong>re</strong> colla<strong>re</strong>d peccaries that <strong>re</strong>qui<strong>re</strong>d in

94 P<strong>re</strong>dation on Neotropical Monkeys
average 240 minutes, so, almost five times mo<strong>re</strong>
than howlers.
<strong>The</strong> p<strong>re</strong>diction that smaller-bodied size
primates a<strong>re</strong> mo<strong>re</strong> vulnerable to p<strong>re</strong>dators than
larger bodied primates does not fit for human
p<strong>re</strong>dators. It has been observed that large
atelines a<strong>re</strong> the p<strong>re</strong>fer<strong>re</strong>d p<strong>re</strong>y for many human
groups, and a<strong>re</strong> systematically selected as
hunting targets (Mena et al., 2000; Alvard &
Kaplan, 1991). Another p<strong>re</strong>diction suggested
that the vulnerability to p<strong>re</strong>dation might be
higher in unfamiliar a<strong>re</strong>as for the primates. Local
human populations tend to know the <strong>monkey</strong>s
home range, and in some cases hunters may
p<strong>re</strong>dict the movements <strong>of</strong> primates populations
through their knowledge <strong>of</strong> fruiting patterns <strong>of</strong>
particular key t<strong>re</strong>e species eaten by <strong>monkey</strong>s or
identifying animal feces in the fo<strong>re</strong>st (e. g. the
Makuna: Århem, 1976). On the other hand, the
p<strong>re</strong>diction that primates a<strong>re</strong> mo<strong>re</strong> vulnerable in
the canopy and fo<strong>re</strong>st edges may apply when being
hunted by human p<strong>re</strong>dators. Based on interviews
in the Venezuelan Guayana and eastern Venezuela,
I <strong>re</strong>ceived information from hunters (Amerindians
and C<strong>re</strong>oles) indicating that hunting pr<strong>of</strong>iciency is
higher when <strong>monkey</strong>s a<strong>re</strong> exposed in the fo<strong>re</strong>st
canopies rather than in dense tangles. <strong>The</strong>y
indicated that it is easier to hunt a spider <strong>monkey</strong>
than tend to be found in the highest canopy than
a capuchin <strong>monkey</strong> sometimes found in tangle
understory fo<strong>re</strong>sts strata or secondary fo<strong>re</strong>sts. In
this sense, Carneiro (1970) indicated that in
Peruvian Amahuaca hunting, success dec<strong>re</strong>ases
with higher dense foliage; however, he <strong>re</strong>ported
that this Amerindian group practiced t<strong>re</strong>e
climbing for hunting howler <strong>monkey</strong>s.
<strong>The</strong> idea that p<strong>re</strong>dation is episodic must be
<strong>re</strong>-evaluated when dealing with human hunters.
Homo hunts primates on a <strong>re</strong>gular basis and one
hunting -p<strong>re</strong>datory- episode by humans might
<strong>re</strong>sult in the killing <strong>of</strong> many members <strong>of</strong> the
<strong>monkey</strong> group. For example, Lizarralde (2002)
<strong>re</strong>ported the case <strong>of</strong> a single hunting day in which
five Barí men <strong>re</strong>turned to the village with 16
spider <strong>monkey</strong>s (Ateles hybridus).
<strong>The</strong> consequences <strong>of</strong> living in group for
avoiding p<strong>re</strong>dation as <strong>re</strong>viewed by Chapman
(2000) must be <strong>re</strong>visited for human p<strong>re</strong>dators.
Contrary to the p<strong>re</strong>diction, living in a large
group is a disadvantage, because <strong>monkey</strong>s may
be mo<strong>re</strong> easily found by hunters. For instance,
practically enti<strong>re</strong> groups <strong>of</strong> woolly <strong>monkey</strong>s (130
individuals) and 11 howler <strong>monkey</strong>s we<strong>re</strong> killed
in one day by a Siona-Secoya hunting party
composed by 286 persons in the Ecuadorian
Amazon (Vickers, 1980). <strong>The</strong> idea that primate
groups might c<strong>re</strong>ate p<strong>re</strong>dator confusions and
the<strong>re</strong>by dec<strong>re</strong>ase the feasibility that each
individual may be p<strong>re</strong>yed while inc<strong>re</strong>asing the
defense against the p<strong>re</strong>dators, does not appears
to function as a potential anti-p<strong>re</strong>datory
adaptation for human hunters. Hunters may kill
the most <strong>of</strong> the whole group or just select
particular individuals based on cultural
p<strong>re</strong>fe<strong>re</strong>nces. In this sense, female primates may
be selected to hunt in order to obtain <strong>of</strong>fspring as
pets. In the case <strong>of</strong> Ateles sp., females may be
selected because they a<strong>re</strong> conside<strong>re</strong>d mo<strong>re</strong> “tasty”
than males (Waimiri-Atroari: Souza-Mazu<strong>re</strong>k et
al., 2000), mo<strong>re</strong> “tasty” than other <strong>monkey</strong> species
such as Saimiri oerstedii (Guaymi: González-
Kirchner & Sainz de la Maza, 1998), or even a<strong>re</strong>
conside<strong>re</strong>d “better” hunting games during the
rainy season because this <strong>monkey</strong> species is fatter
during this t<strong>re</strong>e fruiting period (Matsigenka:
Shepard, 2002).
Boinski & Chapman (1995) provided new
insights on potential di<strong>re</strong>ctions for testing
hypotheses <strong>of</strong> p<strong>re</strong>dation on primates. <strong>The</strong>y
argued that comparisons <strong>of</strong> p<strong>re</strong>dation rates with
group size might <strong>re</strong>p<strong>re</strong>sent a bias because <strong>of</strong> large
intraspecific variability in primate group size.
However, they suggest that other standards for
comparisons like p<strong>re</strong>y interbirth intervals might
be mo<strong>re</strong> fruitful for understanding the effects <strong>of</strong>
p<strong>re</strong>dation on primates. It is important to note
that despite the low f<strong>re</strong>quencies <strong>of</strong> p<strong>re</strong>dation for
both p<strong>re</strong>dators, primates with longer interbirth
intervals such as Ateles sp. and Lagothrix sp. a<strong>re</strong>
the p<strong>re</strong>fer<strong>re</strong>d targets among human hunters (e.
g. Yanomamö: Saffirio & Scaglion, 1982;
Matsigenka: Shepard, 2002; Tabl. 1). This may
<strong>re</strong>sult in low levels <strong>of</strong> primate population
<strong>re</strong>covery and also local extinctions.
Boinski & Chapman (1995: 2) state “on an
evolutionary time scale, inc<strong>re</strong>ased p<strong>re</strong>dation
p<strong>re</strong>ssu<strong>re</strong> may favor large groups, but on a shorter
ecological time scale, high p<strong>re</strong>dation levels may
dec<strong>re</strong>ase group size di<strong>re</strong>ctly, simply through the
death <strong>of</strong> animals.” In this <strong>re</strong>gard, they indicated

that G. B. Stanford studies <strong>of</strong> chimpanzees (Pan
troglodytes) p<strong>re</strong>dation on <strong>re</strong>d colobus <strong>monkey</strong>s
(Procolobus badius) is an instructive example <strong>of</strong> the
effect <strong>of</strong> p<strong>re</strong>dation on an ecological time scale.
Chimpanzees hunting on <strong>re</strong>d colobus (76
observed cases in four years) have <strong>re</strong>duced to
almost the half the size <strong>of</strong> the <strong>monkey</strong> population
that inhabit the chimpanzee groups’ range
compa<strong>re</strong>d to the <strong>re</strong>d colobus populations that live
outside this range (e.g. Gombe site: Stanford,
1998 vs Kibale site: Struhsaker, 1975). In other
words, in the field sites in which Pan density is
higher, Procolobus density is lower (Stanford,
1998).
A similar effect may be occurring in the
Neotropics between human and some <strong>monkey</strong>
populations. Since the P<strong>re</strong>hispanic period,
Amerindians established a close <strong>re</strong>lationship with
non-human primates from cosmological believes
to the thought <strong>of</strong> <strong>monkey</strong>s as food (Baker, 1992,;
van Akke<strong>re</strong>n, 1998; Karadimas, 1999, Braakhuis,
1998, García del Cueto, 1989; Urbani & Gil,
2001; Cormier, 2003). In the p<strong>re</strong>sent work only
Amerindians groups we<strong>re</strong> conside<strong>re</strong>d, assuming
that a<strong>re</strong> the human populations that first ente<strong>re</strong>d
the New World while using long-term traditional
hunting techniques. <strong>The</strong> p<strong>re</strong>decessors <strong>of</strong> cur<strong>re</strong>nt
Amerindian populations occupied the tropical
Americas since circa 11,000 year ago to
approximately 1,250-1,600 A. D., intensely using
a<strong>re</strong>as like lowland Panama and Brazil that have
been historically conside<strong>re</strong>d “pristine” rainfo<strong>re</strong>sts
in the New World (Bush & Colinvaux, 1990;
Colinvaux & Bush, 1991; Heckenberger et al.,
2003). In addition, the<strong>re</strong> is di<strong>re</strong>ct evidence from
an archaeological site in northern Venezuela that
<strong>re</strong>flected potential consumption or used for other
purposes (e. g. pets) <strong>of</strong> <strong>re</strong>d howler <strong>monkey</strong>s from
at least 3,000 years B. P. (Urbani & Gil, 2001). In
principle, these human groups may have
influenced the distribution/survivorship <strong>of</strong> cur<strong>re</strong>nt
primate populations, as might be the case <strong>of</strong>
potential former populations <strong>of</strong> white-faced
capuchin <strong>monkey</strong>s (Cebus capucinus) in the
northern Mayan <strong>re</strong>gion (Baker, 1992). In
addition, MacPhee & Horovitz (2002) suggested
that probably the Pleistocene Antillean <strong>monkey</strong>
Xenothrix mcg<strong>re</strong>gori might be extinct due to
human influence. So, intense hunting p<strong>re</strong>ssu<strong>re</strong>s
over 10,000 years might di<strong>re</strong>ctly influence local
B. Urbani 95
extinctions <strong>of</strong> faunal populations and probably -
still unknown- contributing to shape the structu<strong>re</strong>
<strong>of</strong> cur<strong>re</strong>nt primate populations as might be
tentatively infer<strong>re</strong>d from the <strong>re</strong>sults. <strong>The</strong><strong>re</strong> is
cur<strong>re</strong>nt evidence to support this idea. For instance,
Souza-Mazu<strong>re</strong>k et al. (2000 579; 591) indicated
that among the Waimiri-Atroari in northern
Brazil, “sex ratios <strong>of</strong> spider <strong>monkey</strong>s [Ateles
paniscus] killed we<strong>re</strong> heavily biased towards
females indicating a stronger hunting p<strong>re</strong>ssu<strong>re</strong> on
those individuals.” Thus, it seems that potential
sex-based diffe<strong>re</strong>nces in hunting selection a<strong>re</strong>
affecting the group structu<strong>re</strong> <strong>of</strong> feral Neotropical
primates groups; naturally mo<strong>re</strong> detailed studies
to evaluate the effect <strong>of</strong> selective hunting on the
primate population dynamic a<strong>re</strong> needed.
Mo<strong>re</strong>over, Pe<strong>re</strong>s (1991) described local
extinctions <strong>of</strong> woolly <strong>monkey</strong> (Lagothrix sp.)
populations as a consequence <strong>of</strong> intense hunting
in the Amazonia. He said that woolly <strong>monkey</strong>
population density (number/km 2) varied from 0
and 7 in hunted sites to 17 and 30 in nonhunted
sites. In addition, C<strong>re</strong>ole populations in
Latin America intensively hunt primates for
market networks that expand the limits <strong>of</strong> hunting
from family consumption to hunting for
commercial purposes (Mittermeier, 1987, for
understanding Amazonian Caboblos -Brazilian
C<strong>re</strong>oles- economy: Nugent, 1993). Bush meat is a
p<strong>re</strong>fer<strong>re</strong>d food source in the local marts. For
example, Castro et al. (1975) found that in six
months, in the popular market <strong>of</strong> Iquitos, Peru,
we<strong>re</strong> sold 1,700 Lagothrix sp., 1,396 Cebus sp.,
557 Alouatta sp., 321 Pithecia sp. and 198 Ateles
sp. (approximate calculation by B. U.). Thus,
together with the Congo Basin in Africa whe<strong>re</strong> the
bush meat crisis is agg<strong>re</strong>ssively th<strong>re</strong>atening primate
populations (Peterson, 2003), Amazonia appears
to be the other geographical a<strong>re</strong>a critically th<strong>re</strong>aten
by hunting (Mittermeier, 1987; see Results).
<strong>The</strong>n, to the question, why New World
primates adopt the above indicated antip<strong>re</strong>datory
behaviors even if they a<strong>re</strong> not efficient
against their main p<strong>re</strong>dator, Homo sapiens?. Two
dimensions might be playing a role on it. First,
humans have interacted with New World
<strong>monkey</strong>s during just a short timeframe (~11,000
years) compa<strong>re</strong>d to the long-time period since
primates had been found in the New World; the
late Oligocene when Branisella boliviana

96 P<strong>re</strong>dation on Neotropical Monkeys
inhabited the central part <strong>of</strong> South America
(Kay et al. 2002). On the other hand, human
social practices and material cultu<strong>re</strong> (particularly
weapons) take part in the equation <strong>of</strong> p<strong>re</strong>dation <strong>of</strong>
Neotropical <strong>monkey</strong>s, being factors unique to
Homo in the Neotropics. Nevertheless, extensive
field <strong>re</strong>search is needed to properly add<strong>re</strong>ss this
issue.
Probably one <strong>of</strong> the best ways to inquiry
about the <strong>re</strong>lationship between Homo and New
World primates, including in particular hunting
-p<strong>re</strong>datory- practices, is to look into new
insights from ethnoprimatological perspectives
on the conceptualization <strong>of</strong> <strong>monkey</strong>s by
Amerindians; a <strong>re</strong>search line that have been just
initiated after Sponsel (1997) (in the
Neotropics: Fleck et al., 1999; Cormier, 2002,
2003; Lizarralde, 2002; Shepard, 2002; Urbani
& Gil, 2001). Many Amerindian groups have
special tied <strong>re</strong>lations with <strong>monkey</strong>s, and
associated and classified them as “beings like
humans” (e.g. Kalapalo: Basso, 1973). For
instance, among the Mekranoti, a Tupi language
Amerindian group <strong>re</strong>lated to the Kayapo group,
indicated that other Amerindian groups used the
word “Kaya-po” for <strong>re</strong>ferring them; the word
“Kayapo” means the people that “<strong>re</strong>sembled
<strong>monkey</strong>s” (Werner, 1985: 173). Ethnoprimatological
studies a<strong>re</strong> fundamental for understanding the
<strong>re</strong>lationship between human and non-human
primates. Mo<strong>re</strong>over, these works will be essential
for understanding the past and p<strong>re</strong>sent interconnection
<strong>of</strong> both sympatric primates and even
mo<strong>re</strong>, to comp<strong>re</strong>hend the still unknown but
critical futu<strong>re</strong> <strong>of</strong> them. In addition, at the
moment the<strong>re</strong> a<strong>re</strong> few systematic mentions
describing the New Word primate <strong>re</strong>actions to
human primates, and in principle the<strong>re</strong> a<strong>re</strong>
practically no works specifically describing how
humans have potentially affected social and
grouping behaviors in Neotropical <strong>monkey</strong>s (for
Old World primates: Coppinger & Magui<strong>re</strong>,
1980; Bshary, 2001; Tenaza, 1990; Tenaza &
Tilson, 1985). This is an inte<strong>re</strong>sting a<strong>re</strong>a for futu<strong>re</strong>
primatological <strong>re</strong>search in the tropical Americas.
Mo<strong>re</strong>over, <strong>re</strong>cent data on naïve behaviors and
group composition <strong>of</strong> wild primates (for
chimpanzees: Morgan & Sanz, 2003) may be
fundamental for comparing with primate
populations subjected to <strong>of</strong>ten human contact
(including primatologists), in order to understand
potential diffe<strong>re</strong>nces in primate social structu<strong>re</strong>
due to human p<strong>re</strong>sence or impact.
Acknowledgments
Many thanks to Lo<strong>re</strong>tta A. Cormier, Manuel
Lizarralde, Leslie E. Sponsel and Robert S. Voss for
sending me their publications for a p<strong>re</strong>vious
bibliographic <strong>re</strong>view (Urbani, 2002) that we<strong>re</strong>
useful for thinking in this work. To the UIUC
library staff for make available valuable help in
<strong>re</strong>fe<strong>re</strong>nces searching. To Paul Garber and the
anonymous <strong>re</strong>fe<strong>re</strong>es for their critical suggestions. To
Tania for her passion in the conservation <strong>of</strong> tropical
rainfo<strong>re</strong>sts and her enormous love. B. Urbani is
granted by a Fulbright-OAS Scholarship.
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