Journal of Applied

Evidence for post-depletion sustainability in a mature

Blackwell Publishing, Ltd.

Ecology 2005
42, 460– 468 bushmeat market
GUY COWLISHAW 1, SAMANTHA MENDELSON and
J. MARCUS ROWCLIFFE
Institute of Zoology, Zoological Society of London, Regents Park, London NW1 4RY, UK

Summary
1. The trade in wild animals for meat, or ‘bushmeat’, is perceived as one of the most
important threats to wildlife in the tropics. Unsustainable bushmeat extraction also
threatens the loss of livelihoods. However, the long-term persistence of the bushmeat
trade, documented in Africa over several centuries, suggests that the trade can be sus-
tainable. In this study, we investigate sustainability in a mature bushmeat market in West
Africa (Takoradi, Ghana).
2. Our study, conducted over January and February 2000, combined biological and
socio-economic approaches. Offtake data, including information on species identity,
capture location and sales price, were collected in a market survey. Species biological
data, and the historical price of bushmeat and its substitutes (fish and domestic meat),
were taken from the literature. The theoretical sustainable yield for each species was
estimated using standard algorithms.
3. We tested the hypothesis that the current trade is unsustainable with four predic-
tions: that (1) the number of animals extracted exceeds a theoretical sustainable yield,
(2) larger taxa are depleted more heavily close to the city, (3) the price of bushmeat has
outstripped inflation and (4) the price of alternatives, such as domestic meat and fish,
has fallen relative to the price of bushmeat. None of these predictions were supported.
There was therefore no evidence of unsustainability.
4. Analysis of market profiles and hunter reports suggest that the present pattern of
sustainability is the result of a series of non-random extinctions from historical hunting.
Vulnerable taxa (slow reproducers) have been depleted heavily in the past, so that only
robust taxa (fast reproducers), such as rodents and small antelope, are now traded. These
robust taxa are supplied from a predominantly agricultural landscape around the city.
5. Synthesis and applications. The bushmeat trade can have a severe impact on species
that are vulnerable to overexploitation. However, once these species have disappeared,
the remaining species may be harvested sustainably. Bushmeat management policy
might therefore be improved by adopting a two-pronged approach in which vulnerable
species are protected from hunting, but robust species are allowed to supply a sustainable
trade. The productivity of agricultural landscapes for many bushmeat species indicates
that these areas may play an important role in supporting a sustainable bushmeat trade.
Key-words: bushmeat, extinction, Ghana, hunting, sustainable use
Journal of Applied Ecology (2005) 42, 460–468
doi: 10.1111/j.1365-2664.2005.01046.x

hunting is generally utilitarian in nature (Mace &

Introduction
Overexploitation by human hunters is responsible for
the decline of one-third of all mammal and bird species
threatened with extinction (Hilton-Taylor 2000). Such
© 2005 British Correspondence: Guy Cowlishaw (fax: 020 –7483 2237; e-mail:
Ecological Society guy.cowlishaw@ioz.ac.uk).
Balmford 2000) and is most serious in tropical forests,
where wildlife is harvested for food or ‘bushmeat’
(Bennett et al. 2002; Milner-Gulland, Bennett & SCB
2002 Annual Meeting Wild Meat Group 2003). The
bushmeat trade not only threatens those animal species
that are hunted but also those plant species that depend
on these animals for pollination and seed dispersal
461 (Wright et al . 2000). Consequently, the control of
Sustainability of unsustainable bushmeat hunting is a high priority for
bushmeat markets international conservation. The urgency and import-
ance of achieving this goal is further emphasized by
the role that bushmeat plays in the livelihoods of poor
rural populations across the tropics (Mainka & Trivedi
2002; de Merode, Homewood & Cowlishaw 2004).
Over recent years, a growing number of studies
have described bushmeat harvesting as unsustainable
(e.g. Robinson & Bennett 2000), particularly in Africa
where the volume of extraction is exceptionally high
(Fa, Peres & Meeuwig 2002). The term bushmeat has
therefore become synonymous with overexploitation.
However, the link between bushmeat extraction and
unsustainable use is likely to be more complicated.
African societies have harvested and traded bushmeat
for centuries (e.g. Lewicki 1974; Mendelson, Cowlishaw
& Rowcliffe 2003) and this trade must have possessed
some element of sustainability, at least until recent
times, otherwise bushmeat and all its associated animal
species would have disappeared a long time ago. In
reality, the extent to which the trade is unsustainable is
likely to be variable, contingent on a variety of supply-
and-demand factors such as the available habitat for
bushmeat species and the local human population size.
An improved understanding of this variability in
the bushmeat trade is urgently needed to enhance our
understanding of its impacts and to improve the effi-
cacy of conservation action.
In light of this need, the purpose of the present study
was to investigate sustainability in a mature (long
established) urban bushmeat market in West Africa.
Analysis of such a market should be particularly
insightful because urban markets tend to be associated
with unsustainable exploitation, but a mature market
implies some degree of sustainability.
Our first step was to establish whether the local
bushmeat trade is unsustainable. Sustainability is
notoriously hard to assess in such systems, owing to the
difficulties in reliably monitoring offtake across all prey
species at the necessary spatial and temporal scales.
Consequently, we used a suite of indirect measures.
These do not allow us to prove sustainability outright,
but do provide evidence for or against a lack of sustain-
ability. These measures encompass both biological and
socio-economic data to produce a number of indicators.
Individually these measures are not fully informative,
but cumulatively they indicate whether there is evidence
for unsustainable use.
According to the overexploitation hypothesis, the
current harvesting of species is unsustainable and is
therefore associated with a decline in species popula-
tions and bushmeat availability. Under these conditions,
our indicators should show the following patterns.
1. The observed levels of offtake for each taxa will
© 2005 British
exceed their theoretical sustainable yield (prediction
Ecological Society,
Journal of Applied 1·1).
Ecology, 42, 2. Across taxa there will be a positive correlation
460–468 between body mass and average capture distance from
the city (prediction 1·2). This is a common pattern
associated with unsustainable hunting around popula-
tion centres (Alvard et al. 1997).
3. The market price of bushmeat will outstrip infla-
tion as a result of bushmeat becoming increasingly
scarce (prediction 1·3).
4. The growing scarcity of bushmeat will make it
increasingly more expensive than its alternatives such
as beef, mutton and fish (prediction 1·4). These last two
predictions ensue from both the short-term effects of
reduced supply relative to demand and the long-term
effects of a rise in harvesting costs with increasing
scarcity.
Given that we found no evidence of unsustainable
use, we then investigated those factors that might lead
to sustainability. We hypothesized that the market has
experienced a historical period of overexploitation,
leading to the depletion of the most vulnerable species
and their concomitant disappearance from the market-
place, such that only robust species that can sustain
high levels of offtake remain. Sustainability therefore
arises as a consequence of an ‘extinction filter’
(Balmford 1996). We tested this historical depletion
hypothesis using two predictions: that those species
most vulnerable to overexploitation, i.e. those with the
slowest reproductive rates, would be absent from the
market (prediction 2·1), and that local hunters would
report historical declines but current stability in the
abundance of prey species (prediction 2·2).
Methods
 
Our study focused on the mature urban bushmeat mar-
ket of Sekondi-Takoradi (hereafter Takoradi), Ghana’s
third largest city, located in the Upper Guinea Forest
global biodiversity hotspot (Myers et al. 2000). Tako-
radi has several centuries of recorded settlement and
has grown rapidly over the last 100 years through the
development of gold mining, railways and harbour
facilities. This economic growth has also transformed
the surrounding hinterland into an agricultural farm-
bush matrix, consisting of a mosaic of plantations,
mixed bush fallow (predominantly cocoa, coconut and
oil palm) and remnant tropical forest (including sec-
ondary forest). Bushmeat has been documented as part
of the local diet for centuries (Grubb et al. 1998). Data
were collected in January–February 2000, a period
representative of annual bushmeat trading: it avoided
both the peak hunting season (May–July inclusive:
Holbech 1998) and the closed season (August–November
inclusive), and was described consistently as a typical
month by the traders interviewed.
 
Data were collected using a combination of direct
observation and semistructured interviews (following
462 Magrath 1992), from a representative sample of all
G. Cowlishaw et al. actors in the market (farmer hunters, commercial hunt-
ers, wholesalers, market traders and chopbars). These
data describe a total of 2430 bushmeat transactions
reported by 70 different actors encompassing 16 differ-
ent taxa. Data collected for each transaction included:
taxon identity, weight and condition (fresh/smoked),
identity of purchaser (e.g. wholesaler, the public) and
sale price (in Ghanaian cedis, ¢). For a subset of trans-
actions, additional information included the supplier
of the meat (n = 1745 transactions) and the capture
location (n = 438 transactions between hunters and
market traders). Distances between capture location
and the city market were calculated on the basis of road
distance (the distance travelled by the hunter). Trans-
action weight was either measured directly (n = 1094)
or reported by the seller (n = 1362): both methods
produced the same mean carcass weights and were
combined for this analysis. Supplementary data were
collected on actor perceptions of changes in bushmeat
availability (n = 12 hunters) and the weight, price and
type of fish and domestic meat sold in the market
(n = 1138). All our informants were open and relaxed,
and their responses showed a high degree of consist-
ency when subjected to direct observation and when
cross-referenced to other actors. None of the recorded
trade was illegal, although illegal trade usually takes
place openly where it does occur in Ghana (Ntiamoa-
Baidu 1998). Further information on the Takoradi
bushmeat trade and details of data collection are given
in Cowlishaw et al. (2005) and Mendelson et al. (2003).
 
Our analyses focused on the 10 terrestrial mammals in
the trade comprising 84% of the total biomass sold. All
analyses involving sales price per kilogram were based
on the market value of smoked meat (85% of all retail
sales). Historical data describing 1963 Takoradi mar-
ket prices are taken from Asibey (1966). To investigate
changes in the real value of bushmeat (inflation-
corrected), prices were adjusted according to the Con-
sumer Price Index (CPI) taken from IMF (1980, 2001).
The CPI measures the cost of a standardized basket
of market goods and is the most widely used measure
of inflation. The prices of bushmeat and other types of
meat in 2000 were determined by averaging the mean
sales price across traders. Our expectations about price
changes (predictions 1·3 and 1·4) assume that bushmeat
is a luxury, or superior, good. Otherwise, it would not
become more expensive when scarce because consumers
would switch to less expensive alternatives. Although
we were unable to explore the precise elasticity of bush-
meat consumption in Takoradi (data describing local
household income and expenditure on bushmeat are
© 2005 British
unavailable), our research indicates that bushmeat is a
Ecological Society,
Journal of Applied luxury item. Bushmeat was more expensive per kg than
Ecology, 42, fish, beef or mutton in 1963 (Asibey 1966) and con-
460–468 tinued to be in 2000 (see below), and during our study
consumers in Takoradi preferred to eat bushmeat over
these alternatives when they could afford it (Mendelson
et al. 2003).
Determination of the current annual extraction of
the 10 taxa across the Takoradi catchment took two
steps. In the first step, we calculated species extraction
for the city itself, in three discrete stages. (i) To deter-
mine the number of urban retailers, the number of mar-
ket stalls, nm, was recorded through a complete census
of market places while the number of chopbars, nc, was
estimated by extrapolation from nm and the average per
capita bushmeat biomass purchased by each chopbar
from market traders (Bc, m) and sold by market traders
to each chopbar (Bm,c) (calculated from 187 and 375
independently sampled transactions, respectively).
This extrapolation was based on the assumption that
nc · Bc, m = nm · Bm,c · (ii) To determine the annual urban
sales by these traders, we multiplied nm and nc by the
average number of species carcasses we observed each
selling (per capita) in a typical 1-month period, and
multiplied this by 12. (iii) To estimate annual bushmeat
extraction for all urban sales, we accounted for the
additional informal trade between hunters and con-
sumers, on the basis that such sales may comprise up
to 18% of urban sales (Ntiamoa-Baidu 1998). In the
second step, we calculated total extraction from the
entire catchment in two discrete stages. (i) We first
accounted for rural sales of bushmeat, on the basis that
only 17% of sales by farmer hunters are urban sales
(Falconer 1992). (ii) We then accounted for bushmeat
that hunters eat themselves or give away, on the basis
that farmer hunters sell either a consistent fraction
equal to 69% of all captures (Method 1) (Falconer
1992) or an inconsistent fraction of captures ranging
from 17% to 100% (median 66%) depending on the
species in question (Method 2) (Ntiamoa-Baidu 1998).
These calculations thus produced two estimates of
the annual extraction for each species. These esti-
mates indicate that the formal urban bushmeat sales
(160 000 kg fresh mass) are only 14% of total extrac-
tion (1130 tonnes: mean, Methods 1 and 2) for these
taxa in the Takoradi catchment.
The two estimates of annual extraction for each spe-
cies were compared to the estimated sustainable pro-
duction for each species according to two sustainability
indices (Milner-Gulland & Akçakaya 2001). These
indices calculate a species sustainable production, P,
according to its population density (either at its current
population size, N, or at carrying capacity, K), its
intrinsic rate of population increase, rmax, and its mor-
tality or recovery factor, F. According to the Robinson
and Redford algorithm:
PRR = 0·6K(rmax − 1)FRR
In contrast, the US National Marine Fisheries Service
algorithm states that:
PNMFS = 0·5N(rmax − 1)FNMFS
463 To determine K and N, species population densities,
Sustainability of d, were first taken from Fa & Purvis (1997) or estimated
bushmeat markets through standard allometric relationships (Rowcliffe,
Cowlishaw & Long 2003). The catchment area for
each species was then defined as the area around
Takoradi between its minimum and maximum recorded
capture distances. The landscape around Takoradi
is characterized by a farmbush matrix: a representa-
tive habitat type for all species in the market (habitat
preferences from Grubb et al. 1998). Nevertheless,
species estimates for K and N within the catchment were
set to 0·75d and 0·50d, to account for habitat hetero-
geneity and historical hunting in the matrix, respectively.
To err on the side of caution, we also repeated the
analysis with lower values (K = 0·50d, N = 0·25d ),
but similar results were obtained. Species values of
rmax were also estimated by allometry (Rowcliffe et al.
2003). F RR was set at 0·6, 0·4 and 0·2 for very short-
lived species (rodents, 0·5 – 4 kg), short-lived species
(small ungulates, 4 –14 kg) and long-lived species
(bushbuck, 43 kg), respectively (Robinson 2000); FNMFS
was set at 0·5 throughout (Milner-Gulland & Akçakaya
2001).
These algorithms are commonly used to determine
the maximum sustainable production and thus whether
an observed yield is unsustainable. However, they are
less reliable at pinpointing a sustainable yield because
production is not always maximal (e.g. Robinson 2000).
We therefore made conservative estimates of produc-
tion and sustainability by using modest values of K &
N. Stephens et al. (2002) reported recently that PRR
may overestimate sustainable yields in social species.
However, their conclusions were based on models of
marmots, Marmota marmota, which live in larger and
more complex social groups than any of the species
entering the Takoradi market. In addition, Milner-
Gulland & Akçakaya (2001) found that PNMFS outper-
forms PRR in its ability to correctly predict sustainability.
Here we use PRR because it provides an alternative esti-
mate of sustainable production and because it is also
the most widely used index for estimating sustainability
in the field (thus providing comparability with previous
studies). Nevertheless, it should be noted that the PNMFS
figures are likely to be the more accurate of the two
measures.
Parametric statistical tests were always employed
where the data under investigation did not differ from
a normal distribution (Kolmogorov–Smirnov tests:
P > 0·05). Variables were loge-transformed where this
improved the fit to a normal distribution (e.g. body
mass). Where the data were skewed and could not be
transformed satisfactorily, non-parametric tests were
used. Given the inherent patterns of statistical non-
independence in these data (e.g. one hunter will be
responsible for several transactions, which in turn can
© 2005 British
involve multiple market traders), we used average
Ecological Society,
Journal of Applied species values across all transactions, rather than each
Ecology, 42, individual transaction, as the unit of analysis. All statis-
460–468 tical tests were two-tailed.
Results
  

First, we compared the mean estimates of observed
production (average of Methods 1 and 2) and sustain-
able production (average of P RR and PNMFS) for the
terrestrial mammal taxa in the Takoradi catchment
(Fig. 1). Across species, actual production is consistently
lower than sustainable production (paired-sample t-
test: t9 = −5·0, P = 0·001). On average, observed extrac-
tion is only 20% of what might be sustainable (median,
n = 10 species). In no case does the actual yield exceed
the sustainable yield, although in two cases the actual
yield falls within the range of estimates for the sustain-
able yield: the extraction of these species, Atherurus
africanus and Cricetomys emini, may therefore be at the
limit of what is sustainable. Nevertheless, these results
suggest that the current extraction of bushmeat in the
Takoradi catchment does not exceed a sustainable
harvest, contrary to prediction 1·1.
Secondly, we explored the relationship between taxon
body mass and harvesting distance from Takoradi
(Fig. 2). Contrary to prediction 1·2, there was no sig-
nificant positive relationship between these two varia-
bles, either across all taxa (Pearson correlation: rp =
0·29, n = 10, P = 0·42) or separately across rodents
(rp = −0·12, n = 5, P = 0·84) and ungulates (rp = − 0·82,
n = 5, P = 0·09). There was therefore no evidence of dif-
ferential depletion of larger prey in closer proximity to
the city. Our result was not confounded by smaller spe-
cies being more valuable as the carcasses of larger taxa
always obtained higher prices (rp = 1·00, n = 10, P <
0·001).
Thirdly, the real price of bushmeat (inflation-
controlled) has not increased but rather decreased over the
past 37 years. Contrary to prediction 1·3, the Takoradi
market price of bushmeat has increased by 6052%
between 1963 and 2000, while inflation has increased
by 11 627% in the same period. The real price of bush-
meat has thus declined by nearly one-half (48%) in the
37-year period preceding this study.
Fourthly, contrary to prediction 1·4, a plot of market
prices in 2000 against market prices in 1963 indicates
that the price of bushmeat has also failed to increase in
comparison to the price of other types of meat sold in
Takoradi over this period (Fig. 3). Rather, the price of
bushmeat has declined relative to the price of both mut-
ton and beef, while remaining approximately constant
relative to the price of fish.
   

These results indicate that the current bushmeat trade
may be sustainable. To determine whether this is the
result of historical overexploitation, we identified a set
of potential bushmeat taxa (rodents, primates and
464
G. Cowlishaw et al.

Fig. 1. Comparison between observed and sustainable annual production for terrestrial mammals in the Takoradi catchment.
The observed production (hatched bars) is shown as the mean and range obtained from two alternative calculation procedures
(Methods 1 and 2). The sustainable production is shown for two alternative indices: PNMFS (open circles) and PRR (filled circles).
Species are ordered from top to bottom, first ungulates then rodents, each in order of declining body mass.

Fig. 2. Relationship between species body mass and distance
to market by road from the capture location. Means and
standard errors are shown. Each point is one species: open
circles are rodents, filled circles are ungulates.
ungulates ≥ 700 g body mass) that should occur in the
Takoradi market: each has a previously recorded local
distribution, exists in farmbush matrix habitat and
could be hunted legally during the study. These taxa are
scored for presence/absence in the market and ranked
© 2005 British
by their intrinsic rate of population increase rmax in
Ecological Society,
Journal of Applied Table 1. This analysis indicates that all low-rmax species
Ecology, 42, are absent while all high-rmax species are present. This
460–468 finding supports prediction 2·1.
Fig. 3. The market price of fish, beef and mutton relative to
the price of bushmeat in 1963 and 2000. The position of
commodities above the line indicates the extent to which their
relative price has increased. Means and standard errors are
plotted. Actual prices (¢/kg) are as follows. In 1963 (Asibey
1966): fish = 0·79; beef = 0·82; mutton = 0·99; bushmeat = 1·41.
In 2000 (this study): fish = 4995 ± 12 (n = 5 traders); beef =
6990 ± 18 (n = 5 traders); mutton = 8018 ± 24 (n = 7 traders);
bushmeat = 8533 ± 117 (n = 11 traders). During the study
period, ¢3728 = US$1.
An alternative explanation would be that slow repro-
ducers are naturally rare and their absence reflects the
rarity with which they enter the market. Because the
number of carcasses for each taxon found in the market
can be predicted by its body mass (linear regression:
r2 = 0·31, F1,8 = 5·0, P = 0·056), we can estimate the
465 Table 1. Local farmbush matrix speciesa listed on the Second Scheduleb of the Wildlife Laws of Ghana (Government of Ghana
Sustainability of 1998): primates, ungulates and rodents
bushmeat markets
Massc Presence
Common name Latin name (kg) rmaxd in market

Flying squirrels Anomalurus spp. 0·7 1·09 Present

Pel’s flying squirrel Anomalurus pelii 1·5 0·85 Present
Royal antelope Neotragus pygmaeus 3 0·68 Present
Brush-tailed porcupine Artherurus africanus 3 0·68 Present
Maxwell’s duiker Cephalophus maxwelli 10 0·46 Present
Black duiker Cephalophus niger 20 0·37 Present
Bay duiker Cephalophus dorsalis 20 0·37 Present
Bushbuck Tragelaphus scriptus 42 0·29 Present
Yellow-backed duiker Cephalophus sylvicultor 63 0·26 Absent
Red river hog Potamochoerus porcus 80 0·24 Absent
Giant hog Hylochoerus meinertzhagene 150 0·20 Absent
Lesser spot-nosed monkey Cercopithecus petaurista 4 0·18 Absent
Mona monkey Cercopithecus mona 5 0·17 Absent
African buffalo Syncerus caffer 285 0·16 Absent
Red-capped mangabey Cercocebus torquatus 7 0·15 Absent

a
Species distributions and habitat preferences in Ghana are taken from Grubb et al. (1998). Species listed in order of declining rmax.
b
Species that cannot be hunted during the closed season, but can be hunted as adults at all other times. Three further species occur
in the market but are absent from the Second Schedule because they are hunted without restriction: Atherurus africanus (brush-
tailed porcupine), Cricetomys emini (giant rat) and Thryonomys swinderianus (cane rat, or grasscutter).
c
Species body mass taken from Kingdon (1997).
d
Species intrinsic rate of population increase, rmax, calculated from Rowcliffe et al. (2003).

number of carcasses that should be present for each
absent species on Table 1. In each case, the predicted
number is relatively high (median = 15). Even in the
case of African buffalo, the rarest species, seven car-
casses should have been recorded during the study
period. Across these missing species, the number
expected in the market was significantly greater than
the number observed (i.e. 0) (one-sample t-test: t6 =
3·46, P = 0·013). Thus natural rarity does not appear
to explain the absence of these species.
Finally, of the 12 Takoradi hunters interviewed
about bushmeat species trends, only those who had
been hunting for at least 8 years reported a decline in
prey abundance. In support of prediction 2·2, all hunt-
ers who had been operating for less than 8 years (3–
7 years, median 5 years, n = 4) perceived no change in
abundance, whereas all hunters who had been active
for a longer period (8 –24 years, median 16 years, n = 8)
reported a decline.
Discussion
These analyses suggest that the bushmeat trade in Takoradi
is currently in a sustainable phase. This sustainability
appears to be the result of historical overexploitation
that has eliminated the vulnerable species from the market.
Such ‘extinction filters’ have been widely described in
island ecosystems, where the current fauna comprise
only those species able to survive the anthropogenic
impacts associated with island discovery and coloniza-
© 2005 British
tion (Balmford 1996). Although extinction filters have
Ecological Society,
Journal of Applied not been linked previously to the bushmeat trade, our
Ecology, 42, findings are corroborated by a substantial literature
460–468 on hunter gatherers from both archaeological and
anthropological sources. These studies report a con-
sistent transition towards smaller-bodied (more robust)
prey species in hunter-gatherer harvests over time (e.g.
Cannon 2000; Jerozolimski & Peres 2003). Our results
also highlight the emerging biodiversity crisis in West
Africa, where overhunting has led to the possible
extinction of Miss Waldron’s red colobus monkey Pro-
colobus badius waldroni (Oates et al. 2000; McGraw &
Oates 2002) and widespread local extinctions among
large mammals in Ghana’s national parks (Brashares,
Arcese & Sam 2001).
The pattern of post-depletion sustainability that has
emerged from this study is based on a set of indirect
measures that cannot alone provide conclusive proof
of sustainability. Further research in this area would
therefore be valuable, both in Takoradi and elsewhere.
Additional information that would help to substan-
tiate our findings would include more detailed data on
the abundance and productivity of prey populations in
the Takoradi catchment and the scale of rural extrac-
tion. In addition, further data on temporal changes to
the size of the catchment area and the overall number
of hunters in the system (both of which should remain
constant under stable conditions), and on the costs and
revenues for hunter trips (which should also remain
equal when the system is stable), would be informative
and relatively easy to collect.
It is also important to consider the generality of
these results. Takoradi appears to be a typical Ghanaian
urban market (Cowlishaw et al. 2005), but are there
any unusual aspects of this system that might limit the
application of these results elsewhere? The coastal
location of Takoradi may be important in this respect,
because an abundance of fish in the market might
466 reduce the consumption of bushmeat and thus be
G. Cowlishaw et al. responsible for making the bushmeat trade sustainable
(cf. Brashares et al. 2004). However, bushmeat and fish
are unlikely to be directly interchangeable in Takoradi,
because the former is a luxury good whereas the latter
is not. Moreover, the finding that bushmeat consumption
in Takoradi is 0·01 kg per capita per day (Cowlishaw
et al. 2005), a typical figure for urban dwellers across
Central Africa (Chardonnet et al. 1995), suggests that
consumer demand for bushmeat in Takoradi is charac-
teristic of other cities in tropical Africa.
In addition, similarities between the bushmeat trade
in Takoradi and other localities suggest that the histor-
ical depletion of vulnerable species, followed by an
ongoing trade in more robust species, may be a com-
mon pattern across Ghana. The same five bushmeat
species that comprise 67% of the Takoradi market bio-
mass (cane rat Thryonomys swinderianus, brush-tailed
porcupine Atherurus africanus, Maxwell’s duiker
Cephalophus maxwelli, bushbuck Tragelaphus scriptus
and black duiker Cephalophus niger) also comprise
70% of market biomass across 15 localities in five dif-
ferent Regions (Ntiamoa-Baidu 1998). Similarly, there
is no evidence elsewhere of a scarcity-driven increase in
the price of bushmeat, either in absolute or relative
terms: the real price of bushmeat has declined in
the capital Accra (1975–1993: Tutu, Ntiamoa-Baidu
& Asuming-Brempong 1996), and the price of beef and
mutton have increased more rapidly than the price of
bushmeat in both Accra (1990 – 1993: Tutu et al. 1996)
and Kumasi, Ghana’s second largest city (1980–1986:
Manu 1987). In addition, the pattern of species deple-
tion in Takoradi matches hunter perceptions of species
vulnerability elsewhere in the country: brush-tailed
porcupine A. africanus and Maxwell’s duiker C.
maxwelli are widely thought not to have been affected by
hunting, in contrast to yellow-backed duiker Cephalo-
phus sylvicultor, giant hog Hylochoerus meinertzhagene
and red-capped mangabey Cercocebus torquatus (Western
Region: Holbech 1998), while the largest of the slow
reproducers, giant hog H. meinertzhagene and African
buffalo Syncerus caffer, are also reported to be locally
extinct elsewhere (Ashanti region: Ntiamoa-Baidu 1998).
Finally, evidence that local species depletion is not a
contemporary phenomenon, but rather occurred some
years previously, is also found elsewhere in Ghana.
Falconer (1992) documented that, around Kumasi,
village elders reported a decline in bushmeat species
in their lifetime, but contemporary hunters stated that
prey populations were stable and local traders reported
that bushmeat supply had not declined over the previous
10 years.
 
© 2005 British
These results emphasize the dynamic nature of bush-
Ecological Society,
Journal of Applied meat harvesting systems and the severity of the bush-
Ecology, 42, meat crisis. They also have two important implications
460–468 for conservation policy. First, if mature urban bush-
meat markets are in a sustainable phase, having already
lost their vulnerable species, then those markets should
not be priority targets for conservation action. Rather,
the priority targets should be new markets or those that
are supplied from new catchments (e.g. logging camps:
Auzel & Wilkie 2000). There are perhaps only two cir-
cumstances in which such mature urban markets might
justify significant attention: (1) where a monitoring
programme is desirable, e.g. because a shift is expected
in local socio-economic, demographic or ecological
conditions that might precipitate an elevated demand
for, or reduced supply in, bushmeat; or (2) where con-
servation action is required to assist those species that
are too heavily depleted to appear in the market but still
persist in low-density remnant populations susceptible
to extinction (in this study it has not been possible to
establish whether the depleted species that are absent
from the market are locally extinct or not).
Secondly, and most importantly, if bushmeat mar-
kets can be supplied sustainably solely from robust spe-
cies existing in the farmbush matrix, it is possible in
principle to protect vulnerable species and habitats
without threatening the livelihoods of those people
who depend on the bushmeat trade, many of whom
already live in poverty (e.g. de Merode et al. 2004). Thus
far, the importance of the farmbush matrix as a source
of bushmeat has been largely neglected, with conserva-
tion attention focusing on hunting in low-productivity
primary forests (e.g. Robinson & Bennett 2000). The
farmbush matrix, in contrast, might be much more
productive. This is due partly to the abundance of crops
(Falconer 1992) and partly to the patches of high-
productivity secondary forest (Cowlishaw & Dunbar
2000) in these agricultural mosaics. This productivity,
and also the need to control crop raiders, helps to
explain how bushmeat extraction in secondary forest
can match or exceed that in primary forest (Wilkie
1989) and why Ghanaian hunters prefer hunting in
farmbush matrix over other habitats (Falconer 1992;
Holbech 1998). Moreover, this pattern does not appear
to be unique to tropical Africa. The potential value of
agricultural landscapes for wildlife populations, and
thus for the production of wild meat, has also been
highlighted recently in studies in Costa Rica and Peru
(Daily et al. 2003; Naughton-Treves et al. 2003).
The possibility that city bushmeat markets can be
supplied solely and sustainably by robust species from
an agricultural landscape is an encouraging result. It
suggests that the conservation goals of vulnerable spe-
cies and habitat protection are not always in conflict
with human needs. However, our ability to implement
management steps that allow for the coexistence of vul-
nerable species with an active bushmeat trade will be
more difficult in practice, due to the limited institu-
tional capacity of those countries where bushmeat is
traded (e.g. Smith et al. 2003). In Ghana, wildlife laws
already exist to regulate the bushmeat trade, many of
which are consistent with the policy recommendations
made here, such as the protection of vulnerable species
467 and the permitted hunting of robust species. Neverthe-
Sustainability of less, both public awareness and state enforcement of
bushmeat markets these laws is extremely limited (Mendelson et al. 2003).
Similarly, Rowcliffe et al. (2004) have demonstrated
that the existence of wildlife laws in Democratic
Republic of Congo is in itself insufficient to regulate
bushmeat hunting without effective enforcement. The
development of effective conservation management
on the basis of these research findings is therefore likely
to involve wider issues of capacity building and good
governance (Davies 2002). This remains a significant
challenge for the future.
Acknowledgements
We thank Lars Holbech, Katherine Homewood,
Francis Hurst, Catherine MacKenzie, Candy Mends and
Paul Symonds for their assistance in this research; and
Lars Holbech, E. J. Milner-Gulland, John Robinson,
David Wilkie and two anonymous referees for their
helpful comments on this paper. We are also grateful to
the many actors in the Takoradi bushmeat commodity
chain who generously contributed to the study. The
project was funded by NERC and ESRC. G. C. is
currently in receipt of a NERC Advanced Fellowship.
The fieldwork was carried out in affiliation with the
Protected Area Development Programme of the Wildlife
Department, Ministry of Lands and Forestry, Repub-
lic of Ghana. This paper is a contribution to the ZSL
Institute of Zoology Bushmeat Research Programme.
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Received 19 November 2004 final copy received 9 February 2005
Editor: Rob Freckleton