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Abstract
There are many issues that must be considered when thinking about chimpanzee
conservation, including disease, poaching for bushmeat, and habitat loss. Moreover, there are
strong reasons to justify protecting the species, such as maintaining access to various kinds of
research, preserving the role they play in their ecosystem, and sustaining biodiversity. We
reviewed journal articles to assess the current status of chimpanzees in the wild, the problems
they face, and possible solutions to these problems. Most of the articles we reviewed presented
data indicating that the numbers of chimpanzees are steadily declining. We found habitat loss to
be the primary factor for this decline. It is also the most difficult factor to address. Disease, like
the outbreaks of the Ebola virus that occurred in the mid-nineties, has also ravaged chimpanzee
interaction poses risk for spread of infectious disease if appropriate protocol is not followed.
Finally, while the bushmeat trade is not the most significant cause of decline, it is still an
important contributing factor, and perhaps the one we can most easily attend to. We conclude
that the ways to save the chimpanzee species are complex and vary with the cause of decline, but
Introduction
particular concern is the conservation of our closest relatives in nature, the great apes, and above
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all the chimpanzees (Pan troglodytes). While it is known that chimpanzees are less endangered
than the other great apes (Oates 2006), a survey of the available literature indicates that the
number of chimpanzees surviving in the wild is nonetheless decreasing (Campbell et al. 2008,
Kondgen et al. 2008, Walsh et al. 2003). The well established status of chimpanzees as
humankind’s closest relative in nature makes their conservation a top priority. We share roughly
96% of our genes with them, including 99% of our aligned genes (Varki and Nelson 2007).
Some scientists, such as Jared Diamond, have gone as far as arguing that they should be placed
in the genus Homo. Regardless of one’s position on that more academic matter, their genetic
similarity makes them ideal subjects for medical research (Diamond 2006). Furthermore, their
sophisticated behavior, which includes the ability to use tools and transmit this knowledge
culturally, continues to inform us about the way our own ancestors may have lived (Whiten et al.
2005). Maintaining the role chimpanzees play in their habitats and ecosystems, as well as
preserving biodiversity for its own sake offer us additional incentive to protect them.
The status of the chimpanzee species, and its encompassed subspecies, is one of the key
aspects of its conservation. The species is generally accepted to be made up of four subspecies:
Union for the Conservation of Nature (IUCN) is the worldwide body that authoritatively selects
the status of species and subspecies. The IUCN currently has the species, including all four
subspecies, listed as “Endangered.” This means that over the past three chimpanzee generations
there was a 70% reduction in the population and, if trends continue as they are now, there will be
a 50% reduction in the next three chimpanzee generations (IUCN 2008). This calculation is
performed over generations rather than years because the reduction cannot be seen on a year-to-
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year basis in a productive way. For the current range of the species, as well as the subspecies, see
Figure 1. There is dispute as to what the status of chimpanzees currently is, however. There are
calls to raise the status for the species to “Critically Endangered” (Walsh et al. 2003). There are
also those who think that the status should be dropped to “Vulnerable” (Oates 2006). While the
status at the moment remains in some dispute, the overall population trend for the species has
been agreed upon as decreasing (Walsh et al. 2003; Campbell et al. 2008). For that reason, we
believe it is prudent to err on the side of caution and begin enacting conservation measures now.
Other conflicts in this area occur when one is only concerned with conservation at the
species level. It has been argued that P. t. verus has more genetic variation between it and the
other subspecies than some mammal species, and should therefore be elevated to full species
rank (Morin et al. 1994). The fact that there is so much genetic variation within chimpanzees is a
strong reason for the conservation effort of the species. The possibility that what we currently
think of as chimpanzees may be more accurately described as multiple species would alter the
hominid family tree and possibly change the way that humans look at their own evolution.
In this paper, we will seek to understand the status of chimpanzees and the issues that
affect their risk of extinction. Because the conservation of a species is a complex problem, we
will examine it from many angles. A few of these challenges include the rate of loss of habitat,
the native people’s overexploitation of the species for use in the bushmeat trade, the issue of
disease and its relation between chimpanzees and humans, the scientific conflict over the level of
endangerment of the species, as well as the scientific dilemma about whether certain subspecies
should be elevated to species rank (IUCN/SSC 2003). We will then compare and critique
solutions that various sources put forth. In doing this, we aim to offer an accurate portrayal of
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the chimpanzee’s plight and present any possible solutions. We also hope to justify further
research and action; indeed, there is general consensus that better data are needed to fully
evaluate the state of the chimpanzee and consider appropriate conservation measures.
Methods
To find relevant articles and collect data, we made extensive use of databases such as
Biological Abstracts and Wildlife & Ecology Studies Worldwide to help narrow our focus. We
selected only primary study articles from peer-reviewed scientific journals related to our topic.
The only exception was one review article on the status of the species. As the majority of studies
is necessary to use a paper which combines the individual populations into the total population.
We focused on more recent articles, but we did not have a precise cutoff date. Some older
articles offer valuable data about past conditions like nest counts and ranges that we can compare
with more recent figures. When there were multiple studies about similar aspects of the overall
problem, we compared them critically to see if disagreements of any kind arose, and if so, why
they arose. We also referred to a couple of general texts to verify certain facts and to access
more general background information. As we continue this process, it will be important for us to
focus on whether the articles offer comparable data. This is crucial because it enables us to look
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Ecosystem Impact
The impacts on the ecosystem by chimpanzees are many. One impact that provides
and the red colobus monkey (Colobus badius). This interaction influences behavior for both
species, but there is disagreement as to the degree of interaction, though most of this
disagreement stems from observational methods. Boesch (1994) suggests that the C. badius
populations under study are not habituated to people, and that the presence of human observers is
a confounding factor. Stanford (1995), meanwhile, disagrees and suggests that C. badius
populations in question are habituated to human presence, and that data should be treated
interactions with chimpanzees as C. badius serves as a major food source for chimpanzees in
certain areas. Habituation to humans can affect chimpanzee behavior when it comes to the
hunting of C. badius.
For the Gombe area, where the chimpanzee subspecies P. t. schweinfurthii is found (as
well as red colobus monkey C. b. tephrosceles), Boesch (1994) reports that chimpanzees are
afraid of the mobbing behavior that the monkeys generate in response to a chimpanzee attack.
This leads to a preference by chimpanzees for infant and juvenile C. badius. Stanford (1995)
agrees that there is a preference for infants and juveniles, but states that the chimpanzees do not
exhibit fear when attacking. Although there are these few discrepancies, these two studies are
comparable in the majority of their data. Boesch (1994) also compares a population of
chimpanzees in Côte d’Ivoire (P. t. verus) as well as their corresponding population of red
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colobus monkeys (C. b. badius). Both of these studies indicate that, in the areas where these
species are sympatric, both influence the behavior of the other. As far as C. badius’ part in the
diet of chimpanzees, they are important prey where the ranges overlap (Stanford 1995). An
aspect of the behavior that is discussed is that of vocalizations of C. badius in response to the
presence of chimpanzees. There is dispute over whether the call has been classified or not
(Stanford 1995; Boesch 1994). It is, however, one or two distinct calls that are used only in the
In comparing the Gombe and Côte d’Ivoire subspecies of both species in question, there
are various outstanding differences in behavior. The Côte d’Ivoire monkeys are much nosier, and
thus easier for chimpanzees to locate. Therefore, the chimpanzees in that area actively seek out
the monkeys as prey whereas the Gombe chimps only hunt if they come across a group of
monkeys in the course of other activities (Boesch 1994). Overall, the Côte d’Ivoire chimpanzees
are less fearful of the defenses of the monkeys than the Gombe chimpanzees. The monkeys also
exhibit different behaviors in the two regions, but this appears to mainly be due to the different
tree height and cover (Stanford 1995; Boesch 1994). An aspect of C. badius behavior that has
been agreed upon by multiple sources is that the social structure of the groups (that there are
multiple males) is a direct evolutionary response to the predation on the species by chimpanzees
(Busse 1977; Stanford 1995). It is also evident that the relationship between these two species is
currently evolving as one of the Côte d’Ivoire chimpanzees tried a new strategy of upsetting the
monkeys so that they charge him and he can kill one without the help of his group. This has
worked well for him and he continues this behavior (Boesch 1994). Also, the chimpanzees of
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Gombe, who are generally afraid of the mobbing behavior of the colobus, have a few individuals
that overcome that fear and increase their hunting success (Boesch 1994).
A last critical aspect of these interactions is that the chimpanzees keep the populations of
C. badius below the line at which competition for food becomes an issue that renders their social
groups unstable. This is observed by the fact that when a population of C. badius has its entire
home range within a chimpanzee group’s home range, that group is considerably smaller than a
group that is on the edges of the chimpanzee group’s range (Stanford 1995).
The above explanations of chimpanzee and red colobus monkey interactions show that
chimpanzees are an integral and necessary part of the red colobus ecosystem and must be
Population Estimation
There is overall agreement across most of the literature that the number of chimpanzees
in the wild is declining. A recent survey (conducted from July to December 2007) of
chimpanzee nests in Côte d’Ivoire found a 90% decline compared to a similar survey conducted
from 1989-1991 (Campbell et al. 2008). This study was consistent with another done in other
countries in western equatorial Africa, though this one found only a 56% decline compared with
a 1981-1983 survey of the same area (Walsh et al. 2003). One methodological difference that
could account for this discrepancy is that the second article grouped chimpanzees and gorillas
together, while the first focused exclusively on chimpanzees. While both of these species are
endangered, it is probable that they are not equally endangered which could influence the data.
The study justified grouping the apes together by noting that their nests are hard to distinguish,
which could therefore be a source of error in the survey in Côte d’Ivoire. Additionally, the
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surveys—and previous surveys they were compared to—were carried out at different times and
There is another confounding variable, however, that may be even more important. By
their own admission (and perhaps due to a constraint of resources) the scientists in the second
study surveyed “areas thought to have unusually high large mammal densities” (Walsh et al.
2003). That is, instead of surveying the entire region (as the 1981-1983 survey with which they
compare their data did), they were biased towards specific areas where they expected apes to be.
For this reason, they suggest their estimate for the decline of the ape populations might be
conservative. This may partially explain the rather large gap between the estimates. If this is the
case, it suggests that the more extreme findings from Côte d’Ivoire can be generalized to at least
the rest of this region. Both articles cite human population increase as a primary cause of
chimpanzee population decline and support further research and an elevation of the chimpanzee’s
conservation status to “critically endangered.” The second also mentions the breakout of Ebola
in ape communities. Unsurprisingly, it found that overall the densest ape populations were
located furthest away from urban centers and regions of Ebola outbreak.
However, the status of chimpanzees is not equally dire in all places. High population densities of
chimpanzees and gorillas were observed in a study conducted in northern Cameroon. Using nest
counts, chimpanzee density was estimated at 1.1 per km2 (Dupain et al. 2004). This is in direct
contrast with data from Côte d’Ivoire. In the 2007 survey conducted there, it was estimated that
fewer than 50 individuals inhabited an area of 167 km2. From this estimate, the density can be
calculated at 0.3 chimps per km2 (50/165.5 = 0.3). The discrepancy between densities in
northern Cameroon and Ivory Coast may be partially explained by geography. Chimpanzee
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population densities are generally higher in the Congo Basin—which includes Cameroon—
probably due to the lower human population densities in this region (Tutin 2001).
Yet geography would not account for the low population densities in northeastern Gabon,
which is also part of the Congo Basin. Further, in the Gabon study, the researchers focused on
regions thought to have high large mammal densities, leading them to hypothesize that their
estimate of ape decline might be conservative. Conversely, the Cameroon study was performed
in an area that featured characteristics that seemed unlikely to yield high counts (Dupain et al.
2004). Specifically, the area surveyed was unprotected, inhabited by a tribal community,
routinely attracted hunters, and featured a road and logging company nearby. In response to the
unexpected findings, wherein ape counts in a non-protected area were comparable to those
within the bordering reserve, Dupain et al. advocate incorporating the “matrix,” used to describe
Why Walsh et al. (2003) yielded low population density counts (under favorable
conditions), while Dupain et al. (2004) yielded high population density counts (under
unfavorable conditions) is clearly an issue that must be resolved if we are to better understand
account for these major differences in results. Both studies consisted of counting ape nests, but
Walsh et al. (2003) did not distinguish between gorilla and chimpanzee nests, whereas Dupain et
al. (2004) defined all nests located on the ground as belonging to gorillas and all nests located in
the trees as belonging to chimpanzees. This minor methodological difference should not obscure
the fact that the area surveyed was likely the variable that determined the opposing outcomes.
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Disease
The starkly divergent results of the above studies may be authentic and explainable.
Huijbregts et al. (2003) reported that Northeastern Gabon has been ravaged by the Ebola virus,
suggesting a strong supportive explanation for the seemingly contradictory findings of these
studies. Low population densities were documented in the Minkebe Forest, which is a region
through researchers and eco-tourists (Kondgen et al. 2008). Kondgen et al. maintain that
tourism (whether or not it is educationally based) and research are beneficial to apes, either by
reducing the incidence of poaching or helping to promote the cause of conservation. However,
this close interaction can transmit zoonotic respiratory diseases (Huijbregts et al. 2003;
Woodford et al. 2002). The experiment of Kondgen et al. (2008) involved clinical observations
of infected chimpanzees. In the three communities of chimpanzees studied, morbidity was above
92% and mortality ranged from 3-19%. Necropsies confirmed the presence of pathogens and
phylogenetic analyses suggested a human origin for these (Kondgen et al. 2008).
Formenty et al. (1999) focused specifically on the Ebola virus in Côte d’Ivoire, though
they did not examine it from the context of human transmission. While it seems probable that
chimpanzees could acquire Ebola from humans in theory, in practice the opposite is more likely,
perhaps through the bushmeat trade. Formenty et al. (1999) claim that their study was the first
time an Ebola outbreak was described in nature. It took the form of an epidemiological survey;
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operational definitions such as “probable case,” “definite case,” and “contact” were assigned to
refer to chimpanzees that likely were infected with Ebola, those who certainly were (confirmed
through laboratory testing), and those who came into contact with the infected subjects. Of the
43 chimpanzees in the community, 12 died or were missing (one definite case, seven probable
cases, and four possible cases). Further observation found that casual and even sexual contact
were not high risk factors for becoming infected, but having an infected mother or eating meat
were. Formenty et al. (1999) concluded that due to high mortality (they suggest 100% of those
infected died) and short duration, chimpanzees are not likely to be the source that sustains Ebola.
One implication that follows from this conclusion is that a call to prevent Ebola virus
Gabon (Huijbregts et al. 2003), as the outbreak is possibly one of the major causes of the low ape
population densities in the region described by Walsh et al. (2003). Huijbregts et al. (2003)
utilized reconnaissance walks to execute their survey of the region, a method that has the
advantage of covering a large area relatively quickly, but overlooks certain landmarks, like
swamps and “very dense vegetation”. This could potentially lead the study to underestimate the
number of apes in the region, but the author downplays this, stating that there is a “good
correlation between [reconnaissance] walks and transect data (the typical method) for the same
area” (Huijbregts et al. 2003). Ultimately, only 0.03 chimpanzee nests per kilometer were
observed, compared to 1.88 nests per kilometer at other sites (Huijbregts et al. 2003). The
chimpanzee nest counts are dwarfed not only by counts from other sites, but counts from the
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Minkebe Forest prior to the Ebola outbreaks (these studies occurred before 1994, while the count
being discussed was taken 1998-2000). As the article points out, the most troubling aspect of
these findings is that the Minkebe forest is fairly undisturbed by humans and used to serve as a
The conclusions from the studies concerning Ebola in both Côte d’Ivoire and Gabon’s
Minkebe Forest are consistent and clear. Ebola can have a devastating impact on remaining
chimpanzee populations, and the effect can be even worse in the small, isolated remnant
populations that are increasingly typical of the species as their numbers decline (Huijbregts et al.
2003). That the zoonotic virus can be transmitted between chimpanzees and humans compounds
the problem. Humans are not at fault for the toll Ebola has taken on chimpanzees, but if it was it
not for encroachments in other areas (e.g. deforestation or hunting), the Ebola virus would not
pose such a threat to the species future. Thus, disease is not the primary force behind
the benefits of research and ecotourism while minimizing the spread of infectious disease. After
summarizing the risks of ape/human contact and anticipating many of the concerns raised by
Kondgen et al. (2008), Woodford et al. (2002) offer a set of guidelines that would mitigate
transmission of disease. Generally, these are common sense solutions like screening tourists for
health conditions, reducing proximity, limiting exposure, performing autopsies on deceased apes,
and vaccinations for research personnel and tour guides (Woodford et al. 2002). They also
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facilitate the research of apes, it requires long term close contact and can impact the behavior of
the subjects. For example, habituated gorillas in Rwanda and Uganda began to sleep and forage
outside of their usual ranges, putting them at higher risk for disease, and possibly hunting as well
(Woodford et al. 2002). A similar effect could also occur with chimpanzees, although they may
be more prone to wander out of a home range naturally. This demonstrates the complexity of
2008). Some researchers believe that this threat is even greater than that of habitat fragmentation
(Bowen-Jones E and Pendry S, 1999). The main cause of this overharvest is the bushmeat trade.
Bushmeat is essentially meat that is harvested from wild animals for human consumption (Fa et
al. 2005). The bushmeat trade has major impacts for the species as well as the ecosystem that it
inhabits. Because humans hunt in a way that certain species are preferred, it is typical that larger
animals are harvested sufficient densities that hunters must switch their focus to smaller prey,
usually rodents (Fa et al. 2005). According to the aforementioned sources, the number of
chimpanzees harvested in any part of the species’ range is fairly small when compared to other
species harvested. For example, in a study of the causes of death in the chimpanzee population in
Gombe National Park, only one incident of poaching was recorded; poaching for bushmeat
purposes, however, can make up about 20% of the mortality for a region (Williams et al. 2008).
One of the main reasons why the bushmeat trade can have such a large impact is because
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chimpanzees have a low birth rate. The adults are usually the population segment that is
harvested (Njiforti 1996). This is also the reproductive segment, so reproductive rates drop as a
The average harvest rate of chimpanzees in west and central Africa is about 1.15% per
year (Fa et al. 2005). There are between 172,700 and 299,700 chimpanzees in existence
according to the IUCN (2008). If we assume an average of those two numbers, there would be
236,200 living chimpanzees. Using the 1.15% per year harvest rate, 2716.3 would be killed per
year. One large non-intentional impact on the hunting of this species is the logging industry. The
logging industry is an important source of economic wealth. However, the building of roads
creates new avenues by which hunters can access previously unhunted or poorly hunted areas
Solutions to the problem of chimpanzee harvest for bushmeat are complex. Currently the
preferred method for capturing animals for the bushmeat trade is by using snares largely due to
the overall lower price of cable compared to other harvesting devices (300 African Financial
Community francs per yard vs. 500-2500 francs per gun cartridge) and snares last between two
and three years (Noss 1998). A primary issue is that snaring is currently illegal in the majority of
countries where chimpanzees occur. If snaring were made legal, then there could be efforts to
regulate it that would result is lower catch. The money to regulate it could come from fees to
hunt. One major issue with snaring is that many chimpanzees actually die in the snares. Due to
conditions in these forests, this causes decay and scavenging resulting in a minimum of 27% of
catches being thrown away by the hunter or lost (Bowen-Jones and Pendry 1999, Noss 1998).
One way to reduce this problem would be to legally restrict the distance a hunter can go to hunt
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from the towns and villages that the hunters live in. This is effective because wastage as a result
of those two factors is positively correlated with distance from the dwelling of the hunter
(Bowen-Jones and Pendry 1999, Noss 1998). Legalization would also solve the problem of
confiscating the animals that are captured for bushmeat. By stopping this confiscation, the
hunters would get more money, and therefore need to kill less to obtain the same amount of
money.
A system of quotas is suggested by Bowen-Jones and Pendry (1999) so that the number
removed does not exceed the number produced. This however, is not a very practical system as
there is no one in place to enforce laws such as that. Another more feasible system would be to
get rid of the middleman. Currently the majority of the profits of the commercial sale of
bushmeat (this does not include local sales) go to the middleman (Bowen-Jones and Pendry
1999). Were this individual removed, the hunters would get more money for their sales and
thereby be able to support themselves on fewer animals. The quotas mentioned above would
prevent any overharvesting to get-rich-quick off of the better returns thus generated. Another
aspect that would make the appeal of bushmeat drop is taxation. If bushmeat were to be
legalized, it could be taxed. This would raise its price, thus making it less affordable than the
alternatives such as farm-raised meat. The profits that the government receives from this taxation
program could be returned to the villages of origin in the form of school and health services
Many of these solutions have problems associated with them. The main reason is the
same reason that bushmeat hunting is a problem in the first place, that is to say, lack of
government control. As mentioned above, the great majority of bushmeat hunting is illegal.
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However, it is common for the governments of the countries not have an adequate supply of
money with which to hire rangers and wardens to ensure hunting and trapping laws are followed
correctly. For example, there is a protected area of forest that is 4,000 square kilometers, and
there are only 30 wardens to police it (Noss 1998). This lack of personnel to enforce the laws is
an extreme issue that is more politically than biologically based and creates anger among the
locals. Therefore, the most practical solution is that of expanding the size of protected areas. The
smaller the area of reserve, the more deeply into it hunters will penetrate (Bowen-Jones and
Pendry 1999). That is to say that if there is a reserve of 130 square kilometers, the core of that
park will be more likely to hunted than that of a reserve of 1270 square kilometers. The unhunted
cores of the reserves could then become a source for the hunted populations on their perimeter.
For a visual representation of this, see Figure 2. A solution to the issue of logging in the
bushmeat trade could be making the logging companies put restricted access to their roads so
that only employees and authorized personnel could go through. Also, once the logging operation
is finished, they would have to seal the roads so that they cannot be used again.
Conclusion
a precarious position. While the evidence is not unanimous (the IUCN and its researchers have
the species as declining and “endangered” while Dupain (2004) has the population of their study
as not in danger and not decreasing), most recent nest counts reveal a decline in population
density compared to only twenty years ago. To varying degrees, hunting, habit and disease have
contributed to this decline. Extinction of the chimpanzee would not only be tragic in its own
right, but could potentially destabilize the ecosystems that the species inhabits. Action must be
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taken to curb the downward trend. As the primary cause of the chimpanzee’s plight, humans
have a moral obligation to aid in their conservation. First, elevating chimpanzees to a “critically
endangered” species by the IUCN will mobilize support and resources on the species’ behalf.
Educational programs and ecotourism can help fund further research and raise public awareness,
although the details of the interactions between the chimpanzees and humans must be considered
to prevent transmission of infectious disease. Political solutions must also be pursued especially
in the areas of bushmeat and reserves. Increasing the range of protected regions will limit the
area available to hunters. Legalizing and taxing the bushmeat trade will provide moral and legal
incentives for traders to comply with quotas and provide money to fund and execute other
conservation initiatives. Without urgent and comprehensive action, the chimpanzee’s status will
continue to worsen. Ultimately, both humans and chimpanzees alike will benefit from an
ambitious conservation effort. Scientists will have the continued opportunity to use the
chimpanzee for medical, behavioral, and evolutionary research, and the layperson will continue
to enjoy these gregarious creatures in their natural habitat and zoos alike.
References
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Figures
Figure 1
19
Christopher Warneke
Lab 724 Tuesdays 1:20-4:20
William Gade
Lab 722 Mondays 5:00-8:00
April 21, 2009
The above is the documented range of this species as well as its cousin (the gracile
chimpanzee or bonobo). In this paper we focused exclusively on the robust chimpanzee
(Pan troglodytes). Also, the respective ranges of the various subspecies can be seen. This
figure was reproduced from IUCN/SSC (2003).
Figure 2
20
Christopher Warneke
Lab 724 Tuesdays 1:20-4:20
William Gade
Lab 722 Mondays 5:00-8:00
April 21, 2009
The above figure represents the concept of having a larger reserve to protect a larger core area.
The distance indicated by the two arrows is the distance a hunter will penetrate into any reserve
from any given access point. Due to the fact that there are no roads on these two ideal reserves,
they can only be accessed from the perimeter. As can be seen, the distance of penetration does
not vary with the size of the reserve. The reserve on the left has proportionally more area that is
not hunted than the reserve on the right. Therefore, a greater number of individuals are protected
there. As this is the case, the individuals present would disperse into the hunted area after too
many are in the core area, thus providing a fairly consistent flow of meat to hunters over time,
while the reserve on the right is not as likely to do this as the core area is smaller. This concept
was taken from Bowen-Jones and Pendry (1999).
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