close

Вход

Забыли?

вход по аккаунту

?

Responses of chimpanzees to a recently dead community member at Gombe National Park Tanzania.

код для вставкиСкачать
American Journal of Primatology 74:1–7 (2012)
COMMENTARY
Responses of Chimpanzees to a Recently Dead Community Member at Gombe
National Park, Tanzania
FIONA ANNE STEWART1, ALEXANDER KENNETH PIEL2, AND ROBERT C. O’MALLEY3
1
Department of Biological Anthropology, Leverhulme Centre for Human Evolutionary Studies, University of Cambridge,
Cambridge, United Kingdom
2
Department of Anthropology, University of California-San Diego, La Jolla, California
3
Department of Biological Sciences, University of Southern California, Los Angeles, California
Chimpanzee responses to the death of a group member have rarely been observed in the wild and most
instances involve infant deaths. One of the very few detailed accounts of a group’s response to the death
of an adult community member is from Gombe National Park, Tanzania, where Teleki [Folia
Primatologica 20:81–94, 1973] observed the responses of 16 chimpanzees to an accidental death, none of
whom touched the body. Now, almost 40 years later, we report on the behaviors of 16 (different) Gombe
individuals to the recently dead body of an adult female community member. In stark contrast to
Teleki’s account, we observed individual chimpanzees’ responses to range from curious observation and
passive investigation (e.g. smelling and grooming) to the shaking, dragging, and frustrated beating of
the body. Variation across demographic groups is described and may reflect individuals’ past experience
with death. The implications of our observations are discussed in the context of core morbidity traits
shared between humans and chimpanzees. Am. J. Primatol. 74:1–7, 2012.
r 2011 Wiley Periodicals, Inc.
Key words: chimpanzees; death; gombe; morbidity
INTRODUCTION
Humans exhibit diverse cultural variation in
mortuary practices [Huntington & Metcalf, 1979],
where treatment of dead bodies ranges from their
consumption, to the placing of them as far above or
below ground as possible [Martin, 1996]. Some
treatment patterns, however, are more universal
and consistent with sociobiological explanations.
For example, Littlefield and Rushton [1986] showed
that (across Canada) mothers grieve more intensely
than fathers, and both grieve more intensely for men
than women. Healthy children will be grieved over
more intensely than unhealthy children, and parents
with fewer children will grieve more intensely than
those with many children. Additionally, the type of
death, relationship characteristics between the dead
and the survivor, and the social circumstances
surrounding the death also influence human
responses [Parkes, 1985]. This variation can be observed
in response intensity, duration, or whether the entire
process is immediate or delayed after the death.
Other animals exhibit diverse responses to the
death of conspecifics. Responding to the same
polyamines [Pinel et al., 1981], ants and rats both
remove the dead from their burrows, carrying them
out or burying them, respectively. Dolphins show
both sociosexual behavior and aggression to carcasses [Dudskinski et al., 2003], whereas elephants
consistently express behaviors of ‘‘concern’’ (e.g.
r 2011 Wiley Periodicals, Inc.
monitoring and sniffing the carcass [Douglas-Hamilton
et al., 2006]. Across the primate order, most
examples describe responses to dead infants,
although not all. Over 40 years ago, Cowgill [1972]
described how a pair (alpha male and female) of
captive pottos (Perodicticus potto) began to ration
food for a recently dead (and subsequently removed)
b male cage-mate, seemingly searching the enclosure
regularly for months after the death.
Perhaps no species has seen the history, and more
recent explosion, of reports on behavioral responses to
death as has chimpanzees (Pan troglodytes). Recent
descriptions include accounts on observations from
three research settings: captive [Anderson et al.,
2010], sanctuary [Cronin et al., 2011], and wild [Biro
et al., 2010] populations. These reports build on
previous accounts [Boesch & Boesch-Achermann,
Contract grant sponsor: Jane Goodall Institute.
Additional Supporting Information may be found in the online
version of this article.
Correspondence to: Alexander Kenneth Piel, Department of
Anthropology, University of California-San Diego, La Jolla, CA
92093-0532. E-mail: akpiel@ucsd.edu
Received 8 June 2011; revised 20 July 2011; revision accepted 21
July 2011
DOI 10.1002/ajp.20994
Published online 28 October 2011 in Wiley Online Library (wiley
onlinelibrary.com).
2 / Stewart et al.
2000; Goodall, 1968; Kooriyama, 2009; Matsuzawa,
1997; reviewed in Pettitt, 2011] of chimpanzee
responses to death, almost exclusively to infants and
juveniles. Matsuzawa et al. [1990] describe coming
across the carcass of a 6.5-year-old male with 20
individuals sitting around it in Bossou, Guinea,
although few other data are provided. In the same
community, Biro et al. [2010, p R351] describe the
extended carrying of dead infants by their mothers at
the same site, where individuals of all age-sex classes
‘‘attempted to touch, poke, or handle the bodies,’’ but
not to carry them. Infants were carried for up to 68
days after the infant’s death. In contrast to other
observations from the wild (see below), no aggression
was ever observed toward the carcasses at Bossou.
In one of the rare observations of chimpanzee
responses to the death of an adult, Anderson et al.
[2010] described captive chimpanzee behavior at
Blair Drummond Safari Park, Scotland. Unique to
this report were the detailed observations made from
video before, during, and after the death. The
authors describe behaviors that are analogous to
those that humans exhibit: chimpanzees appeared to
care for the individual before death, test for signs of
life at the point of death, and post-death showed
signs of frustration, and subsequent avoidance of the
location where the death occurred. They proposed
that their observations differed in several ways from
previous accounts. First, while reports from Gombe
[Teleki, 1973] and Tai [Boesch & Boesch-Achermann,
2000] describe excited and aggressive behaviors by
conspecifics around (or in Tai toward) the corpses in
response to death, Anderson et al. [2010, p R350]
observed subdued, calm behaviors, ‘‘strikingly reminiscent of human responses to peaceful death.’’
Additionally, at both Tai and Gombe, chimpanzees
exhibited noticeable interest in the carcass and the
place of death, even dragging the body over short
distances at Tai. The opposite response was seen in
captivity in Scotland, where individuals avoided the
place of death for five consecutive days despite
having slept there the 29 previous nights. The
observations from this small group in captivity
prompted Anderson et al. [2010] to question whether
empathy, self-awareness, and cultural variation—
traits shared by humans and chimpanzees—allow us
also allow a shared awareness of death [see also
Boesch & Boesch-Achermann, 2001].
Finally, a recent report from Chimfunshi Wildlife
Orphanage Trust, Zambia represents one of the first
descriptions that quantifies mother responses to their
dead infants. Although infant carrying is a ubiquitous
behavior across chimpanzee communities, Cronin et al.
[2011, p 420] recognized there is a paucity of evidence
that examines how, in this transitional process, a
mother distances herself from the infant carcass.
Measuring the proximity and interaction between
mother and dead infant across time, the authors
suggest that these types of observed behaviors may
Am. J. Primatol.
specifically inform on ‘‘how chimpanzees respond to the
premature severing of the mother–infant bond.’’
Fashing et al. [2011] detailed the responses of
wild gelada monkeys (Theropithecus gelada) to dead
infants. Females carried dead infants for 1–4 days
before abandoning the body, but geladas in general
exhibited limited or no interest in the death of adult
or juvenile conspecifics. The authors also describe
the death of a mother–infant pair whose physical
weakness eventually prevented them from keeping
up with troop movements. On the first day the pair
failed to follow the troop, group members looked
back often in the direction of the pair, but eventually
moved on to forage. Even upon return to the sleeping
site where the geladas had all slept the night before,
the authors observed no evidence of any attempt to
locate the weakened individuals. The mother died
first, infant at her side. Researchers found the dead
infant alone the following morning [Fashing et al.,
2011]. The authors proposed two explanations for
different responses to death across primates. The first
is ecological, that perhaps extreme (cold or dry)
climates contribute to delayed decomposition of
carcasses, and thus prolonged interest from group
members [Fashing et al., 2011; Matsuzawa, 1997]. The
second is phylogenetic, that these more complex
responses to death (see below) may reflect an ancestral
trait of hominoids only [Warren & Williamson, 2004;
Whilde & Marples, 2011; reviewed in Pettitt, 2011].
Besides that of Anderson et al. [2010], the only
other documented report of chimpanzee responses to
adult deaths comes from Teleki’s [1973] observations
from Gombe. He described in detail the responses of
chimpanzees to the sudden and dramatic death of a
community member, Rix, who fell from a tree and broke
his neck. Behavioral responses varied between individuals, and were hypothesized to vary with age or social
relationship. Some individuals stayed with the corpse
for many hours, but avoided touching it, and almost all
made ‘‘wrah’’ calls, prompting Teleki to conclude that
such vocalizations may have a special significance in the
context of a dead community member.
Here we contribute another case study to this
growing list of observations, describing and quantifying the responses of chimpanzees to a dead adult
female conspecific in natural conditions in Gombe.
Given most sick or wounded animals retreat from
their social group shortly before death [Hart, 1988],
detailed observations from the wild are rare and
difficult to record. We describe here the responses of
18 individuals to a recently deceased adult female,
Malaika, of the Kasekela community. The presence
of many community members allowed our detailed
recording of individual responses and sequential
quantitative analysis of how individuals of different
age and sex classes varied in interest and arousal
around the corpse. Additionally, we explore Pettitt’s
[2011] developmental periods of mortuary phases
(Core; Archaic: Modernizing; Modern; Advanced) in
Responses to a Dead Community Member in Chimpanzees / 3
wild chimpanzee responses to an adult community
member.
METHODS
Study Site and Data Collection
Gombe National Park —35 km2 and the site of
one of the longest continuous field studies of wild
chimpanzees—is situated along the eastern shore of
Lake Tanganyika [Goodall, 1986]. At the time of data
collection, there were three communities in the park:
Mitumba in the north, Kasekela in the center, and
Kalande in the south [Pusey et al., 2007]. All
observations come from the Kasekela community,
which numbered 60 individuals (12 adult, 6 adolescent, 2 juvenile and 6 infant males, and 20 adult,
8 adolescent, 1 juvenile and 5 infant females) at the
time of the observation. Data were collected on a
party of 18 individuals (Fig. 2; 7 females, 10 males,
and 1 infant). The dead individual, Malaika, was
approximately 20 years old and had one known
living offspring, a daughter, Mambo, born into the
Kasekela community in 2004 [Wilson et al., 2005].
Malaika immigrated into the community in 2002, but
remained low ranking and appeared to be socially
peripheral [Murray et al., 2007; O’Malley, pers. obs.].
The research complied with protocols approved by
the Tanzanian National Parks) authority, the Commission on Science and Technology, and Tanzania
Wildlife Research Institute as well as adhered to the
American Society of Primatologists Principles for the
Ethical Treatment of Non Human Primates.
Malaika’s body was discovered on January 5,
2010 in Mkenke Valley by Tanzania National Parks
Chimpanzee Trackers. She had been visibly weak, ill,
and was mostly solitary for several days leading up to
her death (Emma Lantz, pers. comm.), and she was
estimated to have died the previous night or early
that morning based on the absence of flies or signs of
decomposition known to occur rapidly in forest
environments [Boesch & Boesch-Achermann, 2000].
F. A. S. and A. K. P. arrived at 0915 hr to find
Malaika’s corpse and a party of chimpanzees already
present. All behavior near the corpse was documented ad libitum with videography and written notes.
The video camera (Sony Handycam DCR-SR82) was
handheld and remained trained on the body to
capture all interactions and activities in close
proximity to it. Videography was interrupted
between 1116 hr and 1230 hr due to low battery
warnings and resumed until 1300 hr. For the entire
observation period, all individuals present, their
behavior, and in particular any interest and each
physical contact with the body were recorded ad
libitumusing written notes and later verified against
the video footage. Individuals were identified on
sight by National Parks Chimpanzee Trackers and
Gombe Research Assistants and on video later
confirmed by RCO. Owing to the difficulty of
simultaneously recording behaviors of all the individuals present, observations focused on individuals
interacting with the body directly or who were
within approximately 20 m of it.
Following Teleki [1973], and in order to directly
compare frequency of behaviors, total time was
segmented into 15-min intervals to assess the
presence and frequency of behaviors. The number
and identity of individuals observed to perform any
of six categories of behavior during each 15-min
interval were used for analyses. Categories of
behavior included investigate, play, vocalize, rest,
feed, and display. ‘‘Investigate’’ refers to behaviors
directed toward the corpse, and as defined by Teleki
included the behaviors ‘‘watch’’ and ‘‘sniff’’ only. We
have included a wide range of behaviors, which were
grouped into five sub-categories describing magnitude of investigation, ranging from passive to
intensive: watch, sniff, poke genitals and sniff
fingers, groom, gentle-handle, rough-handle. Gentlehandle included touch, stroke, hold hand, or embrace
body. Rough-handle included shake, hit, slap,
stamp on, and drag body. Following the removal
of the body, ‘‘investigate’’ refers to behaviors
directed toward the last spot where the body
had lain. The number of 15-min intervals within
which an individual was observed to perform each
of these behaviors was used for analyses of variation
with individual age. In order to control for an
individual presence, the number of 15-min intervals
in which an individual was present within observation range was also recorded, and the proportion
of 15-min intervals in which investigative behaviors
occurred was calculated for each individual.
RESULTS
From 0915 until 1227 hr, 18 individuals
interacted to varying degrees with Malaika’s body
(Figs. 1 and 2). At 0915 hr, her body lay prone on the
ground near a dry streambed while a juvenile
slapped her body on the ground (Movie ESM1) and
five individuals watched silently from arboreal
locations. One night nest was nearby, and although
unknown if this was Malaika’s from the previous
night, the absence of other nests in the vicinity
suggests that most of the chimpanzees arrived at the
site that morning. The alpha male, Ferdinand,
displayed with the body at 0928 hr, throwing her
down into the streambed. This display was one of
several by different males. Malaika was visited by a
succession of individuals who one by one, briefly
investigated the body, while other individuals continued to watch from above. Periodically young
males dragged her body. Soon thereafter, eight
individuals formed a tight circle around the
body beneath a thicket in silence. Most were only
looking, but a few sniffed or groomed the body.
During the observation period, her body was also
Am. J. Primatol.
4 / Stewart et al.
Fig. 1. Number of individuals who performed behaviors during 15-min intervals. The black line indicates the time of removal of
Malaika’s body, after which "Investigate" occurs toward the body’s last location. (Individual ages are from Wilson et al. [2005]).
Fig. 2. Number of 15-min intervals individuals, grouped by sex and age [Teleki, 1973], were present (black line) and were observed to
watch, sniff, groom, poke genitals, and sniff fingers, gentle-handle, or rough-handle Malaika’s body (stacked bars).
sequentially monopolized by individual young males,
which displaced younger chimpanzees. Persistent
rough-handling of Malaika’s body by these young
males suggests that they may have been frustrated
with her lack of response, or else were using her body
as a tool in displaying to community members
[Teleki, 1973]. Malaika’s body was never left alone,
and was eventually dragged over 60 m by at least
four individuals. This dragging (Movie ESM 2)
sometimes appeared as attempts to monopolize
access, as individuals dragged the body away from
others, while at other times, as attempts to haul the
corpse with the party after other individuals had
begun to travel.
The number and identity of individuals who
performed any of six categories of behavior during
15-min intervals are shown in Figure 1. Feed, rest,
and play were infrequent, although the slight
increase in rest over time may indicate a gradual
loss of interest in the carcass. Displays toward and
around the corpse were also infrequent in contrast to
previous accounts [Teleki, 1973]. ‘‘Waa’’ and ‘‘wraa’’
Am. J. Primatol.
calls were the most distinctive of frequent vocalizations, suggesting high levels of arousal and distress.
The most notable difference between previous
accounts and ours concerns levels and type of
interaction with the corpse. Teleki [1973] reported
low levels of investigation (never more than two
individuals in any 15-min period), whereas up to
eight individuals inspected or manipulated the body
in any 15-minute period during our observations.
Teleki also observed no tactile inspection,
whereas we saw a wide range of exploratory acts,
ranging from passive (watch) to intensive (roughhandle—Fig. 2), more similar to observations made
by Boesch and Boesch-Achermann [2000]. These
‘‘passive’’ forms are part of the natural chimpanzee
behavioral repertoire, while the ‘‘intensive’’ are
unusual for any within-group context, wild or
captive. Individuals of all ages and both sexes
intently watched the body, and except for three
females and two males, all sniffed it. No females
except Malaika’s daughter, Mambo, handled the
body. Males rough-handled the body significantly
Responses to a Dead Community Member in Chimpanzees / 5
more frequently than females (Mann–Whitney
U test, U 5 14, n1 5 7, n2 5 10, P 5 0.016, two-tailed),
and younger males more than older males (Spearmans,
r 5 0.731, n 5 10, P 5 0.016, two-tailed). Younger
males also groomed the body and tended to gentlehandle her more than older males (r 5 0.787,
n 5 10, P 5 0.007, and r 5 0.57, n 5 10, P 5 0.085,
two-tailed). These sex-differences and correlations
with male age remain significant if controlled for the
amount of time spent within the vicinity (roughhandle sex difference: U 5 14, P 5 0.016; roughhandle vs. male age: r 5 0.71, P 5 0.022; groom vs.
male age, r 5 0.77, P 5 0.009), with the exception of
rough-handle which then does not correlate with
male age (r 5 0.53, P 5 0.12).
Malaika’s body was removed by Gombe
researchers at 12:38 hr, after all chimpanzees had
moved away from the body. However, some individuals probably remained 40 m away and watched
the body being packaged and removed. After it was
removed, eight individuals immediately returned to
the last location of the corpse. Five individuals
inspected where the corpse had lain by touching
and sniffing the ground. Fudge was the first to
approach, arboreally, and looked down to where the
body had lain. He began a pant hoot that was joined
in chorus by the individuals present. This is reflected
in the peak of the number of individuals vocalizing
during these two 15-min intervals (Fig. 1), of which
vocalizations included pant hoot chorus, wraa call,
and whimper (wraa and whimper indicating distress). Mambo was the last to inspect the site, while
Sparrow and Ferdinand waited a few meters away
before the party finally departed at 1300 hr.
DISCUSSION
Attitudes Toward Death
These observations lend themselves to comparisons from previous accounts. Hosaka et al. [2000]
discuss the diversity of ways chimpanzees respond
to the death of community members at Mahale.
They suggested that the level of decomposition and
cause of death influence emotional state, ranging
from fear to curiosity, with females and juveniles
tending toward the former and males the latter.
Those emotions are then expressed through
vocalizations, most notably in ‘‘wraa’’ or ‘‘huu’’
calls. Mahale males were often aggressive with
carcasses, using them in display behaviors. It is
noteworthy that chimpanzees exhibit similar behaviors also to the carcasses of other animals, while
other primates seem to pay little attention to such
carcasses [e.g. vervet monkeys, Struhsaker, 1967].
This suggests a ‘‘broad curiosity’’ by chimpanzees
whereby they are constantly learning about their
surroundings [Cronin et al., 2011; Hosaka et al.,
2000].
Cronin et al. [2011] described chimpanzees’
investigation of a dead infant’s hands and face as
providing tactile, olfactory and potentially gustatory
signs, possibly to assess the current state of the body.
At Tai, Boesch and Boesch-Achermann [2000]
describe how chimpanzees often lick the blood of
chimpanzees that have been injured, but not from
those recently dead, thus differentiating between the
two states. Teleki [1973] suggested that ‘‘the
behavior of certain individuals [y] seemed to
indicate awareness of a change from activity to
inactivity of a group member, but it remained
uncertain whether any participant grasped the
conceptual difference between life and death’’ and
Pettitt’s [2011] interpretation of Tai chimpanzees’
responses to the death of a 10-year-old juvenile
(Tina)—which included unusual vocalizations, sitting silently, grooming, displaying, and dragging her
body–is that ‘‘something very different is occurring
at the death site.’’ That is, individuals exhibit a
range of behaviors, otherwise not seen in a single
context. He concluded ‘‘chimpanzees appear to be
aware that death has occurred.’’ The passivity of
adult chimpanzees to Malaika’s body could also
suggest an understanding of Malaika’s change of
state, but quantitative comparison of changes in
behavior toward an individual pre- and post-death
are necessary to test these hypotheses. The younger
individuals’ persistent and aggressive physical
manipulation of the body suggested frustration and
possible exaggerated attempts to arouse Malaika.
Anderson et al. [2010] also suggested this type of
inspection and male aggression toward the corpse
may reflect attempts to resuscitate the body. Alternatively, these behaviors could be interpreted as
curiosity at a nonresponsive conspecific.
What can we learn from this diverse behavioral
repertoire in chimpanzees to the death of community
members? Littlefield and Rushton [1986] showed
that variation in type and intensity of response to
death in humans is influenced by the sex of the dead
and griever, as well as the type of death, circumstances surrounding the event, and the nature of the
relationship between griever and the deceased. As
Malaika was an immigrant and relatively lowranking [Murray et al., 2007], she may not have
had strong relationships to any nonkin in Kasekela.
As might be predicted, then, Mambo’s behavior
differed from all others. She watched more continuously, from closer distance, and was the only female
to handle Malaika. She visited the carcass frequently, but briefly as she was often displaced or
distracted by play. Mambo also participated in nine
of ten observed play bouts.
To Littlefield and Rushton’s list, we propose that
we should add the age of the griever, at least for
chimpanzees. Our observations showed that females
and older males interacted less with the corpse.
Sparrow’s high level of interest may result from her
Am. J. Primatol.
6 / Stewart et al.
son’s (Sinbad) persistent handling of Malaika.
Eowyn, a juvenile female, that watched only, is
known to have experienced her mother’s death
(M. Wilson, pers. comm.). One possibility is that
older individuals have previously experienced death,
thus exhibiting less interest. Alternatively, younger
individuals may exhibit also more exploratory behaviors across a range of contexts if they have generally
higher levels of curiosity and lower levels of
neophobia. That there is variation in responses to
death across age and sex classes provides a possible
avenue for testing these hypotheses using pre- and
post-death comparisons of individual behavior in the
wild (if the opportunity unfortunately arises).
Implications for Early Hominid Morbidity
The behavior of chimpanzees should not be
misinterpreted as that of the last common ancestor
of great apes and humans. Nonetheless, in a similar
way that contemporary human behavior at funerals
includes ritual practices that are context-specific
(regardless of variation across cultures), so might a
context-specific behavioral repertoire accompany
chimpanzees in response to the death of a community member. Similarities across these two closely
related species may reveal shared ancestral tendencies. Specifically, Pettitt [2011] has argued that
observations of other primates can help us to identify
core behaviors toward the dead that may reflect our
evolutionary history. These include, but are not
restricted to
1. Cronus compulsions—the urge to dismember,
injure, or consume parts of bodies of conspecifics;
2. Manifestations of morbidity (As defined by Pettitt
[2011, p 8] as ‘‘an enquiring concern with the
injured, diseased, or dead body, whether or not
this derives from a desire to understand the
nature or cause of death of an individual.’’)—
especially by those with a close relationship to the
corpse, in the form of grooming usually;
3. Manifestations of mourning—here possibly
expressed in distress vocalizations (e.g. ‘‘wraa’’
barks), carrying the corpse as a sign of detachment;
4. Social theater around the corpse, including controlled access to the body.
Our observations, like some previous [e.g.
Anderson et al., 2010], of the responses to Malaika’s
death include examples of all of these. (This includes,
according to Pettitt [2011], controlled access of the
corpse, sometimes using it in displays. Behaviors
here are not observed in other contexts and can also
be termed ‘‘funerary gatherings.’’)
Although historically there have been few
reports of primates responding to the death of group
members, this trend is changing, and the last 2 years
Am. J. Primatol.
especially have seen a wave of descriptions, both
from captive and wild settings, and to infants and
adults alike. Combined, these observations not only
shape what Anderson [2011] proposes to be a
‘‘primatological perspective’’ on death, but also
reveal intra- and interspecific similarities in the
treatment of conspecific dead bodies. These patterns,
in turn, lend themselves to reconstructing the
origins of human responses to death.
Uncontrolled observations of spontaneous
events in nature preclude conclusive explanations
but allow comparative (opportunistic) systematic
research to be pursued in future. Studies of both
individual and group-level variation in primate
responses to death contribute to the growing field
of primate thanatology [Anderson et al., 2010] and a
more comprehensive understanding of the evolutionary roots of mortuary practices in the homininae
[Pettitt, 2011].
ACKNOWLEDGMENTS
We thank Gombe Stream Research Centre staff
for long-term data collection and research funded by
the Jane Goodall Institute. We thank the Commission for Science and Technology, Tanzania Wildlife
Research Institute (F.A.S., A.K.P., R.C.O.) and
Tanzania National Parks (R.C.O.) for research
permission; W.C. McGrew, J.J. Moore, J.R. Anderson,
C. Murray, A. Pusey, E. Lonsdorf, M. Wilson, and
B. Hahn for comments and discussion.
REFERENCES
Anderson JR. 2011. A primatological perspective on death.
American Journal of Primatology 73:410–414.
Anderson JR, Gillies A, Lock LC. 2010. Pan thanatology.
Current Biology 20:R349–R351.
Biro D, Humle T, Koops K, Souse C, Hayashi M, Matsuzawa T.
2010. Chimpanzee mothers at Bossou, Guinea carry the
mummified remains of their dead infants. Current Biology
20:R351–R352.
Boesch C, Boesch-Achermann H. 2000. The chimpanzees of
the Tai forest: behavioral ecology and evolution. Oxford:
University Press. 316 p.
Cowgill UW. 1972. Death in Perodicticus. Primates
13:251–256.
Cronin KA, van Leeuwen EJC, Mulenga IC, Bodamer MD.
2011. Behavioral response of a chimpanzee mother toward
her dead infant. American Journal of Primatology
73:415–421.
Douglas-Hamilton I, Bhalla S, Wittemyer G, Vollrath F. 2006.
Behavioural reactions of elephants towards a dying and
deceased matriarch. Applied Animal Behaviour Science
100:87–102.
Dudskinski KM, Sakai M, Masaki K, Kogi K, Hishi T,
Kurimoto M. 2003. Behavioural observations of bottlenose
dolphins towards two dead conspecifics. Aquatic Mammals
29:108–116.
Fashing PJ, Nguyen N, Barry TS, Goodale CB, Burke RJ,
Jones SCZ, Kerby JT, Lee LM, Nurmi NO, Venkataraman VV.
2011. Death among geladas (Theropithecus gelada): a broader
perspective on mummified infants and primate thanatology.
American Journal of Primatology 73:405–409.
Responses to a Dead Community Member in Chimpanzees / 7
Goodall J. 1968. The behaviour of free-living chimpanzees in
the Gombe Stream Reserve. Animal Behavior Monographs
1:165–311.
Goodall J. 1986. The chimpanzees of Gombe: patterns
of behaviour. Cambridge: Harvard University Press.
673 p.
Hart BL. 1988. Biological basis of the behavior of sick
animals. Neuroscience and Biobehavioral Reviews 12:
123–137.
Hosaka K, Matsumoto-Oda A, Huffman M, Kawanaka K.
2000. Reactions to dead bodies of conspecifics by wild
chimpanzees in the Mahale Mountains, Tanzania. Primate
Research 1:1–15.
Huntington R, Metcalf P. 1979. Celebrations of death: the
anthropology of mortuary ritual. Cambridge: Cambridge
University Press.
Kooriyama T. 2009. The death of a newborn chimpanzee at
Mahale: reactions of its mother and other individuals to the
body. Pan Africa News 16:19–21.
Littlefield CH, Rushton JP. 1986. When a child dies—the
sociobiology of bereavement. Journal of Personality and
Social Psychology 51:797–802.
Martin DP. 1996. On the cultural ecology of sky burial on the
Himalayan Plateau. East and West 46:353–370.
Matsuzawa T. 1997. The death of an infant chimpanzee at
Bossou, Guinea. Pan Africa News 4:4–6
Matsuzawa T, Sakura O, Kimura T, Hamada Y, Sugiyama Y.
1990. Case-report on the death of a wild chimpanzee (Pan
troglodytes verus). Primates 31:635–641.
Murray CM, Mane SV, Pusey AE. 2007. Dominance rank
influences female space use in wild chimpanzees, Pan
troglodytes: towards an ideal despotic distribution. Animal
Behaviour 74:1795–1804
Parkes CM. 1985. Bereavement. British Journal of Psychiatry
146:11–17.
Pettitt P. 2011. The paleolithic origins of burial. London &
New York: Routledge.
Pinel JPJ, Gorzalka BB, Ladak F. 1981. Cadaverine and
putrescine initiate the burial of dead conspecifics by rats.
Physiology and Behavior 27:819–824.
Pusey AE, Pintea L, Wilson ML, Kamenya S, Goodall J. 2007.
The contribution of long-term research at Gombe National
Park to chimpanzee conservation. Conservation Biology
21:623–634.
Struhsaker T. 1967. Ecology of vervet monkeys (Cercopithecus
aethiops) in Masai-Amboseli Game Reserve, Kenya. Ecology
48:891–904.
Teleki G. 1973. Group response to accidental; death of a
chimpanze in Gombe National Park, Tanzania. Folia
Primatologica 20:81–94.
Warren Y, Williamson EA. 2004. Transport of dead infant
mountain gorillas by mothers and unrelated females. Zoo
Biology 23:375–378.
Whilde J, Marples N. 2011. The behavior of a zoo-housed
infant orangutan after the death of its mother. Zoo Biology
30:205–211.
Wilson ML, Collins A, Wallauer WR, Kamenya S. 2005. Gombe
stream research centre annual report. Unpublished.
Am. J. Primatol.
Документ
Категория
Без категории
Просмотров
3
Размер файла
144 Кб
Теги
recently, gombo, community, parks, response, chimpanzee, tanzania, dead, national, members
1/--страниц
Пожаловаться на содержимое документа