Ecological, Behavioral, and Life-History Correlates of

Mammal Extinctions in West Africa

JUSTIN S. BRASHARES
Centre for Biodiversity Research, University of British Columbia, 6270 University Boulevard, Vancouver, BC V6T
1Z4, Canada, email justinb@zoology.ubc.ca

Abstract: Identifying the biological traits of species that predispose them to extinction is a focus of research
in evolutionary ecology and conservation biology. This research has traditionally been divided between
studies of extinction or decline in undisturbed habitat islands and studies of the persistence of species af-
fected adversely by human influence. I combined these approaches to test for correlations between nine eco-
logical, behavioral, and life-history traits and vulnerability to local extinction for 41 species of carnivores,
primates, and ungulates in fragmented and exploited habitats in Ghana, West Africa, while accounting sta-
tistically for phylogeny. Species distributed in isolated populations were most prone to local extinction, and
monogamous species and those wherein males defended small harems were also prone to extinction. Body
size, fecundity, abundance, habitat specialization, trophic group, and the degree to which hunters and con-
sumers preferred a species generally were unrelated to species persistence. Although population isolation
and mating system were the only traits that explained a significant amount of the observed variation in
persistence of all species, analyses of carnivores, primates, and ungulates as groups yielded varied results.
Mammals most prone to local extinction in my study reserves were also those listed by the World Conserva-
tion Union as being at greatest risk of global extinction. Thus, my results suggest that the relative isolation
of populations and the mating system displayed by mammals may be good general predictors of their per-
sistence.

Correlaciones Ecológicas, Conductuales y de Historia de Vida de Extinciones de Mamíferos en África Occidental

Resumen: Un tema de investigación en ecología evolutiva y biología de la conservación es la identificación
de las características biológicas de las especies que las predisponen a la extinción. Esta investigación tradicion-
almente se ha dividido entre estudios de extinción o declinación en islas de hábitat no perturbado y estudios
de la persistencia de especies afectadas adversamente por la influencia humana. Combiné estas aproxima-
ciones para probar correlaciones entre nueve características ecológicas, conductuales y de historia de vida y
la vulnerabilidad a la extinción local de 41 especies de carnívoros, primates y ungulados en hábitats frag-
mentados y explotados en Ghana, África Occidental, considerando la filogenia estadísticamente. Las especies
distribuidas en poblaciones aisladas fueron más susceptibles a la extinción local, y especies monógamas y es-
pecies en las que los machos defienden pequeños harems también fueron susceptibles a la extinción. El
tamaño del cuerpo, la fecundidad, la densidad poblacional, la especialización de hábitat, el grupo trófico y el
grado en el que cazadores y consumidores prefieren una especie generalmente no se relacionaron con la per-
sistencia de la especie. Aunque el aislamiento de la población y el sistema de apareamiento fueron las únicas
características que explicaron una cantidad significativa de la variación observada en la persistencia de to-
das las especies, el análisis de carnívoros, primates y ungulados como grupos produjeron resultados varia-
dos. Los mamíferos más susceptibles a la extinción local en las reservas estudiadas fueron los enumerados
como en mayor riesgo de extinción global por la World Conservation Union. Por lo tanto, mis resultados
sugieren que el relativo aislamiento de poblaciones y el sistema de apareamiento mostrado por mamíferos
pueden ser buenos predictores generales de su persistencia.

Paper submitted December 4, 2001; revised manuscript accepted July 15, 2002.
733

Conservation Biology, Pages 733–743

Volume 17, No. 3, June 2003
734 Correlates of Extinction in West Africa
Introduction
Ecologists and conservation biologists have long sought
to identify the ecological and life-history traits that ren-
der species vulnerable to extinction (reviewed in
McKinney 1997 ). Identifying traits associated with ex-
tinction may facilitate the protection of species sensitive
to disturbance. Previous work on traits related to extinc-
tion have focused primarily on the persistence of popu-
lations following habitat fragmentation (e.g., Blake
1991; Laurance 1991; Newmark 1991; Terborgh et al.
1997; Davies et al. 2000) and on prehistoric species loss
as a result of habitat loss and isolation ( e.g., Diamond
1984; Richman et al. 1988; Burkey 1995; Foufopoulos &
Ives 1999 ). This previous work documents extinctions
thought to have been primarily “natural” as a result of
demographic and environmental stochasticity and isola-
tion effects, rather than a result of direct disturbance by
humans.
Although human hunting and collecting have affected
the composition of ecological communities for thou-
sands of years, the last 200 years have brought unprece-
dented rates of extinction from harvest (Meffe & Carroll
1997; Robinson & Bennett 2000 ). Much of the world’s
biodiversity now occurs in habitats fragmented and used
by humans (Whitmore & Sayer 1992; Kramer et al. 1997;
Robinson et al. 1999), but some species in these habitats
tolerate harvest and habitat fragmentation better than
others (Diamond 1984; FitzGibbon et al. 1995). By iden-
tifying traits of species that make them susceptible to
harvest and habitat fragmentation, we may be able to
better target our management efforts to forestall de-
clines in the future (Pimm 1991).
I tested for correlations between nine ecological, be-
havioral, and life-history traits and vulnerability to local
extinction for 41 species of large mammals in six nature
reserves in Ghana, West Africa. Nearly 30 years of cen-
sus counts for these reserves document 78 local extinc-
tions of carnivores, primates, and ungulates with no sub-
sequent recolonization (Brashares et al. 2001). Reserve
size and human population density around reserves
were the strongest correlates of extinction rates in
Ghana ( Brashares et al. 2001 ). My goal was to identify
traits of species correlated with their susceptibility to lo-
cal extinction, after the effects of reserve size on persis-
tence had been accounted for statistically. I also sought
to identify the best models for conservation practice
( Draper & Smith 1998 ) based on the traits that predict
extinction.
The threat to biological diversity in West Africa is high
relative to other areas of Africa because of high human
population density, hunting, and habitat loss (Sayer et al.
1992; Struhsaker & Oates 1995; van Schaik et al. 1997).
Thus, it is worth asking whether results based on the
local extinction of mammals in West Africa can help
wildlife managers identify extinction-prone species in
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Volume 17, No. 3, June 2003
Brashares
other parts of Africa and the world. Therefore, I asked
whether the species found to be most prone to extinc-
tion in Ghana are also considered most threatened with
global extinction by the World Conservation Union (IUCN)
(IUCN 2000).
The nine traits of species I considered for analysis
were population isolation, harem size, home range size,
body mass, habitat specialization, abundance, fecundity,
trophic group, and human preference ( raw data for
traits and dependent variables are available from the
author ). Species with large body size, low abundance,
low fecundity, specialized habitat requirements, and iso-
lated populations are thought to be most vulnerable to
extinction ( Pimm 1991; Case et al. 1992; Rosenzweig
1995; McKinney 1997; Davies et al. 2000). These predic-
tions derive mainly from “natural” extinctions resulting
from demographic and environmental stochasticity and
isolation effects. Attempts to model the persistence of
terrestrial vertebrates under heavy hunting have also
identified mating system, home range size, trophic level,
and human preference as predictors of vulnerability to
extinction (Bodmer et al. 1997; FitzGibbon 1998; Greene
et al. 1998; Woodroffe & Ginsberg 2000).
Methods
Dependent Variable: Species Persistence
Approximately monthly, the Ghana Wildlife Division
conducts counts of all large mammals along 10- to 15-km
foot patrols around ranger posts in each of Ghana’s re-
serves. These game counts began soon after the creation
of Ghana’s protected-area system in the late 1960s, and
they continue today. I limited analyses to large mammals
( 2 kg) counted between 1969 and 1998 in at least two
of six reserves in Ghana. The six reserves range in size
from 58 to 4840 km2, and all occur in savanna habitat
(see map in Brashares et al. 2001).
I used count data to estimate the persistence time, or
time to local extinction within each reserve, for each
species known to be present in a reserve between 1968
and 1970. Only species recorded on five or more counts
during the first 3 years following park establishment
were counted as among those present in a reserve’s ini-
tial complement of species. Species were assumed to
have become locally extinct in the last year they were
observed. Thus, a species recorded 20 times annually in
a park between 1970 and 1980, but not observed after
1980 was assigned a persistence time of 10 years. Spe-
cies judged as initially present in a reserve and observed
at least once from 1996 to 1998 were deemed extant
and assigned a persistence time equal to the age of that
reserve. When time gaps of 6–16 months existed be-
tween observations of a species, I assumed that species
remained extant, rather than becoming extinct and later
Brashares
recolonizing. In practice, there was no gap in observa-
tions larger than 16 months, and 90% of gaps were 6
months or less. Final assessments of species presence
and extinction were corroborated against the species ac-
counts in 27 government reports and seven publications
( references in Brashares et al. 2001 ). The final data set
included 158 occurrences of 41 different species. At the
outset of censuses, 12 species occurred in all six reserves;
5 species occurred in only two reserves.
To identify traits of species correlated with their prob-
ability of extinction, I performed two types of analyses.
In the first set of analyses I compared the mean persis-
tence of species across all reserves against the nine spe-
cies traits. To control statistically for the potential influ-
ence of reserve size on species persistence, I first
included all 158 species occurrences in a linear regres-
sion of log persistence time (dependent) against reserve
size (independent). I then used the residuals of this re-
gression (Fig. 1) to calculate a mean residual persistence
time for each of 41 species. Mean residual persistence,
hereafter called residual persistence, was the dependent
variable used in the first set of analyses. In the second
set of analyses, I compared persistence time against spe-
cies traits separately for each of Ghana’s six savanna re-
serves.
Independent Variables: Species Traits
To determine isolation of mammal populations, I esti-
mated for each species the distance between each re-
serve in which the species occurred and the nearest
known population of conspecifics. The estimates were
based on published and unpublished accounts of wildlife
populations in protected and unprotected areas at approxi-
mately the time the reserves were gazetted (Brashares et
al. 2001).
I used published data to calculate a mean harem size
for each species. This value estimates the mean number
of adult females defended by one reproductively suc-
cessful adult male. I did not account for bachelor or sat-
ellite males or subdominant male herd/pack members
that are prevented from breeding. I used data for West
African populations when possible.
A survey of published literature provided several esti-
mates of home-range size for each of the species I con-
sidered. I calculated a mean value for each species and
used this value in analyses. I did not differentiate exclu-
sive and overlapping home ranges and, when possible, I
used data from West Africa.
For each species, I took the average of body masses
provided for both males and females in several pub-
lished sources ( see Brashares et al. 2001 ). When possi-
ble, I relied on measurements specific to West African
subspecies.
To estimate degree of habitat specialization, I used a
habitat-specialization index ( HSI ) that represented the
Correlates of Extinction in West Africa 735
Figure 1. Species persistence times in six reserves in
Ghana against reserve size. Number of species in re-
serves is 20, 30, 22, 27, 31, and 28, from smallest re-
serve to largest. Persistence and reserve size were posi-
tively related (n  158; r  0.38; p  0.001).
total number of distinct habitat types in which a species
occurs. First, following Kingdon (1997) and Estes (1991),
I identified nine basic African habitat types: desert, sub-
desert/semidesert, dry bush/scrub, dry savanna, wet sa-
vanna, moist/mixed woodland, forest mosaic, lowland
forest, and Afromontane. I then determined the habitat
preferences of species using published accounts. I in-
cluded in each species assessment habitats in which the
species commonly occurs or has occurred within the
last 100 years. The HSI values for species considered in
this study ranged from the minimum of 1 to 8 out of 9.
I estimated the initial abundance of species by sum-
ming the total number of observations of each species in
a reserve during the first 3 years of wildlife counts. This
resulted in 158 separate estimates of abundance, or one
for each occurrence of a species in a reserve. For analyses
across reserves, I derived a single relative estimate of
abundance for each of the 41 species by averaging the re-
siduals from a regression of log10 abundance (dependent)
against reserve size (independent). This produced a sin-
gle estimate of mean residual abundance, hereafter
called mean abundance, for each of 41 species. For
within-reserve analyses, abundance estimates in reserves
were unaltered and are hereafter called abundance.
I used published records of longevity corrected for bi-
ases ( see below ), age at first breeding, interbirth inter-
val, and average litter size to calculate an estimate of
maximum fecundity for each species (Cole 1954). Lon-
gevity records for 21 species were limited to accounts of
captive animals, and information on both wild and cap-
tive populations was available for the rest. Where both
types of information were available, the longevity of spe-
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Volume 17, No. 3, June 2003
736 Correlates of Extinction in West Africa Brashares

cies in captivity was on average 40% greater than that of

wild populations. However, this difference was greater
for carnivores ( 50% ) than for ungulates ( 40% ) and pri-
mates (30%). Therefore, for the 21 species for which no
field data were available, I reduced the reproductive
span reported for captive populations by 30–50%, de-
pending on the species group.
I assigned species to one of three trophic groups: carni-
vores, herbivores, and omnivores, following Estes (1991)
and Kingdon (1997).
To score human preference for each species, I calcu-
lated the economic value of species from market surveys
and combined these data with preference rankings ob-
tained from the literature ( Asibey 1974; Fa et al. 1995;
Juste et al. 1995; Njiforti 1996) and through interviews
with 34 hunters in Ghana. I then placed species into one
of four preference categories based on their cumulative
score: 1, most preferred; 2, preferred; 3, not preferred;
4, least preferred.

Accounting for Phylogenetic Relatedness

To identify traits correlated with species extinction in
Ghana’s reserves, I used Felsenstein’s (1985) method of
phylogenetically independent contrasts to recalculate
the dependent variables, “residual persistence” and
“persistence,” and all nine trait values (PDAP program of
Garland et al. 1993; reviews in Harvey & Pagel 1991;
Martins & Garland 1991 ). The final result of Felsen-
stein’s (1985) independent-contrasts method is, in prin-
ciple, a phylogenetically independent and identically dis-
tributed data set consisting of N  1 standardized
contrasts for each variable ( Felsenstein 1985 ). These
corrected data sets can then be analyzed with conven-
tional regression analysis but with regression forced
through the origin (Garland et al. 1992).
The hypothesized phylogenetic tree I used to calcu-
late independent contrasts represents a consensus of
molecular and paleontological information (Fig. 2). The
hypothesized branching patterns of major clades are
based on those of Gatesy et al. (1999) and Waddell et al.
(1999). Other branching patterns and estimates of diver-
gence times are based on those of Purvis (1995), Arna-
son et al. ( 1999 ), Bininda-Emonds et al. ( 1999 ), Penny
et al. (1999), and Brashares et al. (2000).
Correlating Traits and Species Persistence
I used correlation analysis to identify statistical associa-
tions between trait values and the persistence of all 41
species and to evaluate potential collinearity and inter-
dependence of species traits. Correlation analysis was
performed on both the phylogenetically corrected and
uncorrected data. Values of isolation, mating system,
Figure 2. Hypothesized phylogenetic relationships and
estimated divergence times of 41 species of mammals
included in analyses. Branch lengths and branching
pattern represent a consensus of molecular and pale-
ontological data from seven sources (see methods).
home range size, and body mass were log10-transformed
for analyses to satisfy assumptions of normality.
To identify “best” models, the persistence values of 41
species of mammals were regressed against the nine
traits with forward and backward step-wise analyses on
the phylogenetically corrected and uncorrected data for
comparisons both across and within reserves. I chose best-
fitting regression models using standard statistical mod-
eling methods (Mallows Cp and Akaike information cri-
teria [AIC] statistics; Draper & Smith 1998 ), with entry
of traits into the model set at   0.05 and exit at  
0.10. I then repeated this analysis independently for car-
nivores, primates, and ungulates for comparisons across
reserves. Two of these mammal groups were monotypic
with regard to trophic group, so this trait was not in-
cluded in this level of analysis. For comparisons across
reserves, the persistence values of species were weighted
in regression analyses by the number of parks in which
they occurred ( 1/√n ). Details of techniques for includ-
ing weightings in regressions of phylogenetically cor-
rected characters have been given by Bonine and Garland

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Brashares Correlates of Extinction in West Africa 737

Table 1. Correlation coefficients among phylogentically corrected values of residual species persistence and ecological, behavioral, and life-
history traits for all species ( n  40).a

Trophic Harem Home range

Body mass Abundance Isolation HSI b Fecundity group size size Preference
Persistence 0.095 0.100 0.642*** 0.110 0.195 0.080 0.354* 0.320* 0.139
Body mass 0.186 0.324* 0.395* 0.012 0.416** 0.351* 0.685*** 0.417**
Abundance 0.056 0.028 0.035 0.437** 0.197 0.062 0.178
Isolation 0.376* 0.280 0.084 0.135 0.480** 0.072
HSIb 0.233 0.035 0.225 0.265 0.173
Fecundity 0.138 0.016 0.019 0.385*
Trophic group 0.258 0.087 0.236
Harem size 0.225 0.296
Home range size 0.492***
a
Significance: *p  0.05; **p  0.01; ***p  0.001.
b
Habitat-specialization index.

(1999). I used SYSTAT 9.0 (SPSS 1998) for all regression
and correlation analyses.
Correlating Species Persistence in Ghana with Their
Conservation Status
To evaluate the wider applicability of the best-fitting
models from this study, I used correlation analysis to test
for a relationship between the residual persistence of
mammals in Ghana and IUCN (2000) assessments of the
vulnerability of the same species to global extinction.
The IUCN ( 2000 ) assessments were converted to five
ordinal values for statistical analyses: 1, critically endan-
gered; 2, endangered; 3, vulnerable; 4, conservation de-
pendent; 5, least concern.
combined ( n  40) and for primates and ungulates inde-
pendently, but it was not included in the best model for
carnivores (Table 2).
Species in which males defended large harems were
less prone to extinction than monogamous or weakly
polygynous species ( n  40, p  0.03; Fig. 3 ). Harem
size was related positively to residual persistence in un-
gulates ( n  23, p  0.003) but not in carnivores ( n 
8, p  0.58) or primates ( n  9, p  0.12). Harem size
was also included in the best-fitting models for all spe-
cies combined ( n  40) and for ungulates ( n  23; Ta-
ble 2).
Species with small home-range sizes persisted longer
than species with large home ranges ( n  40, p  0.04;
Fig. 3). This relationship was particularly evident for car-
nivores ( n  8, p  0.007) but was weak or absent for

Results
Table 2. Best-fitting models from forward and backward step-wise
Traits Correlated with Species Persistence across Reserves multiple regression of residual persistence from analyses of
phylogentically corrected data.a
Correlations among phylogenetically corrected values of
Variable Coefficient p
residual persistence, isolation, harem size, body mass,
habitat specialization, home range size, abundance, fe- All species ( n  40) b

log isolation 0.512 0.001

cundity, trophic group, and human preference were of-
log harem size 0.053 0.028
ten significant, but never exceeded 0.69 (Table 1). Ungulates (n  23)c
The best-fitting models identified isolation and harem log isolation 0.170 0.005
size as the best overall predictors of persistence across log harem size 0.244 0.002
reserves (Table 2). Correlations among traits chosen in log body mass 0.064 0.036
Primates (n  9)d
the best-fitting models were not significant and never
log isolation 1.801 0.013
exceeded 0.40. Overall, conventional regression analy- habitat specialization 0.129 0.054
ses yielded results similar to those from analyses using Carnivores (n  8)e
phylogenetically independent contrasts. log home range size 0.540 0.001
Species with populations in close proximity to one an- log body mass 0.520 0.043
a
other persisted significantly longer than species that Best models were chosen with Mallows Cp and Akaike information
were more isolated ( n  40; p  0.001; Fig. 3). Popula- criteria (AIC) statistics (Draper & Smith 1998). The dependent vari-
able is persistence time of species averaged across all reserves in
tion isolation was also correlated with residual persis- which they occurred. Corrected values were of all ecological, life-his-
tence for carnivores ( n  8, p  0.02), primates ( n  9, tory, and behavioral traits.
p  0.02), and ungulates ( n  23, p  0.03). Isolation
b
Best model for all species, adjusted R2  0.56.
c
Best model for ungulates, adjusted R2  0.62.
was selected in step-wise multiple regression as a signifi- d
Best model for primates, adjusted R2  0.64.
cant variable in the best-fitting models for all species e
Best model for carnivores, adjusted R2  0.92.

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738 Correlates of Extinction in West Africa Brashares

Figure 3. Traits of species plotted against persistence in reserves for eight carnivore species (open circles), nine pri-
mate species ( filled circles), and 24 ungulate species (squares). Trend lines are the best-fitting straight lines according
to least-squares regression. Associated statistics are provided in Table 1. The plots shown represent uncorrected data.

primates (n  9, p  0.87) and ungulates (n  23, p 
0.10). Home range size was selected as the best predic-
tor of local extinction of carnivores in a step-wise multi-
ple-regression analysis (Table 2), but it was not selected
in similar analyses for all species or for primates or ungu-
lates alone.
Body mass was unrelated to residual persistence for all
species considered together (n  40, p  0.56; Fig. 3 )
and for carnivores (n  8, p  0.41), primates (n  9, p 
0.93 ), and ungulates ( n  23, p  0.10 ) considered
alone. Body mass was not selected by forward or back-
ward step-wise regression analysis in the best-fitting
model for the complete data set or in models for primates
alone. However, body mass was included in the best-fit-
ting models for carnivores and for ungulates (Table 2).
In general, large mammals that occur naturally in few
habitat types ( i.e., high habitat specialization ) were as
likely as species that occur in many habitats to become
locally extinct (n  40, p  0.50; Fig. 3). There was no
effect of habitat specialization for carnivores (n  8, p 
0.63) or ungulates (n  23, p  0.72) considered inde-
pendently. However, primates classified as habitat spe-
cialists were more likely than generalists to become lo-
cally extinct ( n  9, p  0.05 ). Habitat specialization
was also included in the best-fitting model for primates
(Table 2).
Species of large mammals that occurred at low mean
abundance at the time Ghana’s reserves were gazetted
were not more likely than abundant species to become
extinct subsequently (n  40, p  0.54). Similarly, there
was no significant relationship between mean abundance
and species persistence for carnivores (n  8, p  0.53),
primates (n  9, p  0.45), or ungulates (n  23, p 
0.13). Mean abundance was not selected as a significant
variable in the best-fitting model for the complete data set
or in models for the three mammal groups.
Fecundity was unrelated to residual species persis-
tence (n  40, p  0.23). This was also true in indepen-
dent analyses of carnivores (n  8, p  0.56), primates
(n  9, p  0.22), and ungulates (n  23, p  0.83). Fe-
cundity was also not selected in best-fitting regression
models of all mammals or for the three mammal groups.
Species at the top of the food chain (i.e., carnivores)
were as likely to become locally extinct as species at
lower trophic levels (n  40, p  0.63; Fig. 4). Trophic
group was not included in the best-fitting model for all

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Brashares Correlates of Extinction in West Africa 739

Figure 4. Traits of species related to their persistence in reserves in Ghana. Box plots show the median (horizontal
line), interquartile range (box), and range of the data (whiskers). Associated statistics are provided in Table 1.
The box plots shown represent uncorrected data.

mammals, and I did not consider this trait in regression size. My results were similar with species persistence av-
analyses of individual mammal groups. eraged across reserves and with reserves considered in-
The economic value of species in regional markets dependently, and they were unaffected by statistical cor-
and hunter’s prey preferences (i.e., human preference) rections for the relatedness of species. Correlations
were also unrelated to the residual persistence of mam- among traits chosen in the best-fitting models for both
mals in reserves ( n  40, p  0.39; Fig. 4 ). This result across- and within-reserve analyses were not significant,
was true also for primates (n  9, p  0.76) and ungu- so regression analysis of these traits is not likely to be con-
lates (n  23, p  0.09). I found little evidence of varia- founded by multicollinearity of the independent variables
tion in preference for carnivores and excluded this (Neter et al. 1996; Draper & Smith 1998). I will now con-
group from analysis. Human preference was not se-
lected in any best-fitting model.
Table 3. Best-fitting multiple-regression models from analyses of
phylogentically corrected data for each of six savanna reserves.*
Traits Correlated with Species’ Persistence within Reserves
Variable Coefficient p
Overall, the traits chosen as the best predictors of persis-
Mole National Park (n  28)
tence in regressions were similar among reserves and
log harem size 0.065 0.000
similar to the traits selected in the analyses of all species habitat specialization 0.009 0.013
across reserves (Table 3). For example, similar to across- log isolation 0.020 0.039
reserve results, harem size was selected as a significant Digya National Park (n  31)
predictor of persistence for five of the six reserves, and log isolation 0.157 0.000
log harem size 0.039 0.005
population isolation was selected for four reserves.
log body mass 0.107 0.044
Bui National Park (n  27)
Persistence in Ghana as it Relates to Risk of log isolation 0.174 0.000
log harem size 0.137 0.018
Species Extinction Gbele Resource Reserve (n  22)
log harem size 0.256 0.002
I found a strong relationship between the persistence of
log home range size 0.715 0.024
mammal populations within reserves in Ghana and the Kalakpa Resource Reserve (n  30)
risk of species (i.e., global) extinction as assessed by the log isolation 0.138 0.000
IUCN ( 2000 ). Mammals prone to local extinction in Shai Hills Resource Reserve (n  20)
Ghana also tended to be those at risk of extinction globally log abundance 0.180 0.005
log harem size 0.088 0.008
(rank correlation: n  41, rs  0.85, p  0.001; Fig. 5).
*Best models were chosen with Mallows Cp and Akaike information
criteria (AIC) statistics (Draper & Smith 1998). The dependent vari-
able is persistence time of species. N represents large-mammal rich-
Discussion ness in 1968–1971. Best model for reserves: Mole National Park, ad-
justed R2  0.53; Digya National Park, adjusted R2  0.59; Bui
National Park, adjusted R2  0.71; Gbele Resource Reserve, adjusted
For the 41 species studied, persistence was related nega- R2  0.70; Kalakpa Resource Reserve, adjusted R2  0.43; Shai Hills
tively to population isolation and positively to harem Resource Reserve, adjusted R2  0.67.

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740 Correlates of Extinction in West Africa
Figure 5. Persistence of species in reserves in Ghana
and their global conservation status as assessed by the
World Conservation Union (2000). The solid line is
the best-fitting straight line according to least-squares
regression (n  41; rs  0.85; p  0.001).
sider my results for each of nine traits in light of previous
work and discuss some of their general implications.
Isolation was the best correlate of persistence in this
study, consistent with theory ( MacArthur & Wilson
1967; Levins 1969; Hanski 1991 ) and empirical studies
(Kattan et al. 1994; Stouffer & Bierregaard 1995; Davies
et al. 2000). The persistence of species in overhunted or
other “sink” habitats may depend on continued immigra-
tion from productive source habitats (Roff 1974; Hilborn
1979; Novaro et al. 2000). In Ghana persistence was high-
est in the largest and most remote reserves (Brashares et al.
2001), and it is likely that animals occasionally disperse
from these relatively remote areas to sink habitats else-
where. Populations in reserves isolated from immigra-
tion probably experience a higher risk of extinction as a
consequence (Hanski 1994).
If isolation limits immigration, we should expect that
species that disperse over long distances would be less
susceptible to isolation on average ( Hanski 1994 ). To
test this idea, I derived a second estimate of isolation
corrected for dispersal ability as the ratio of isolation dis-
tance (see Methods) to average dispersal distance (raw
data available from the author; Sutherland et al. 2000). The
ratio of isolation and dispersal distance was correlated neg-
atively with persistence (n  40, r  0.61, p  0.001),
consistent with the expectation mentioned above.
Monogamous to weakly polygynous species suffered
higher rates of local extinction than moderately to
highly polygynous species. This correlation existed
across and within reserves and was particularly strong
among ungulates ( Tables 2 & 3; Fig. 3 ). Greene et al.
( 1998 ) and Legendre et al. (1999) predicted that popu-
lations of monogamous and weakly polygynous species
will be particularly vulnerable to changes in the propor-
tion of breeding females caused by demographic sto-
chasticity or hunting and consequently will be most
Conservation Biology
Volume 17, No. 3, June 2003
Brashares
prone to extinction. My results are consistent with these
predictions, but monogamy might also be related to ex-
tinction because of its link to other traits. McKinney (1997)
invoked this “multicollinearity” argument to explain vari-
ous relationships among related traits, but it seems an un-
likely explanation here. Harem size was unrelated to popu-
lation isolation and home range size, which were each
related to persistence. Harem size was also unrelated to
abundance, fecundity, and habitat specialization, which I
expected might predict persistence (Table 1).
Mammals with larger home ranges were more prone
to local extinction than species with smaller ranges, es-
pecially in the case of carnivores (Fig. 3). This finding is
consistent with other studies of carnivore persistence in
fragmented and heavily hunted habitats ( e.g., FitzGib-
bon 1998; Woodroffe & Ginsberg 2000 ). Home-range
size may be a particularly good predictor of persistence
for carnivores because their large home range size
makes them liable to move outside protected areas and
come into conflict with humans (Woodroffe & Ginsberg
2000). Home-range size was a weaker predictor of per-
sistence for ungulates and primates (Fig. 3).
Body mass was unrelated to persistence in each re-
serve and all reserves combined (Table 3). This finding
is contrary to theory ( see McKinney 1997 ) but consis-
tent with results from empirical studies that have shown
that body size is only occasionally related to extinction
in mammals, birds, reptiles, fishes, or insects (Soulé et
al. 1988; Laurance 1991; Case et al. 1992; Angermeier
1995; Davies et al. 2000). It is possible that differences
in the taxonomic scale and traits I considered greatly af-
fect the strength of correlations between body size and
extinction ( Blackburn & Gaston 1994 ). Indeed, body
size often is linked to population isolation, abundance,
home range size or other traits that better predict ex-
tinction rate (McKinney 1997).
For species in heavily hunted habitats, the expected
relationship between body size and persistence remains
unclear. Where hunters prefer large prey, it seems rea-
sonable that large-bodied species will be vulnerable to
extinction when faced with harvest. Despite this expec-
tation, however, few researchers having looked for rela-
tionships between body size and persistence in hunted
habitats, including me, have found one ( e.g., Bodmer
1995; Fa et al. 1995; FitzGibbon et al. 1995). In practice,
the species most desired by hunters often comprises
only a small percentage of the species extracted, which
may explain why smaller and larger-bodied species have
similar extinction rates (FitzGibbon 1998).
In contrast to previous studies (Diamond 1984; Pimm
et al. 1988; Foufopoulos & Ives 1999), I found that spe-
cies that occurred naturally in fewer habitat types per-
sisted as long in reserves as those with broader habitat
preferences. Several species, including elephant ( Lox-
odonta africana), wild dog (Lycaon pictus), and aard-
vark (Orycteropus afer), are habitat generalists that per-
Brashares
sisted poorly in reserves in Ghana. In contrast, however,
primates that occurred in few habitat types were extir-
pated more often than habitat generalists (Table 2). Spe-
cifically, habitat-restricted colobid monkeys ( Colobus
sp. and Procolobus verus ) persisted poorly compared
with habitat generalists such as baboons (Papio anubis)
and patas and green monkeys (Cercopithecus patas and
C. aethiops).
The relative abundance of individual species was a
poor predictor of their persistence in Ghana. This result
is contrary to theory (Leigh 1975; Goodman 1987; Pimm
1991 ) and empirical results ( Foufopoulos & Ives 1999;
Davies et al. 2000 ) that suggest rare species and small
populations are prone to extinction in isolated and frag-
mented habitats. However, the expectation that rarity
predicts persistence is based on the assumption that ex-
tinction occurs as a result of “natural” demographic pro-
cesses, rather than as a result of human influence, such
as hunting. History provides many examples of how ex-
ploitation by humans has affected species abundances
and how human preference for particular prey has re-
sulted in the local and global extinction of formerly abun-
dant species (Vermeij 1993; Rosenzweig 1995). Further-
more, if hunters in Ghana forage optimally ( Charnov
1976; Alvard 1998 ), they might be expected to maxi-
mize their return by harvesting the most abundant prey
species.
Fecundity and persistence were unrelated in my study,
contrary to the expectation that species with low repro-
ductive rates are vulnerable to extinction in heavily
hunted landscapes ( Bodmer 1995; Bodmer et al. 1997;
Novaro et al. 2000). Fecundity was also unrelated to the
abundance of species in reserves, which suggests that
my estimates of fecundity are a poor reflection of the
productivity of mammals in Ghana. It is likely that many
factors, including resource availability, competition, and
predation, play a larger role than fecundity in regulating
the natural abundance of mammals in Ghana.
Species at the top of the food chain showed rates of
extinction similar to those at lower trophic levels (Fig. 4).
This result contrasts with the notion that carnivores are
among the first to suffer the deleterious effects of habi-
tat fragmentation and hunting ( Schonewald-Cox 1983;
Belovsky 1987 ). Larger-bodied carnivores such as lions
(Panthera leo), spotted hyenas (Crocuta crocuta), and
wild dogs were all prone to local extinction in Ghana.
The vulnerability of these species was in contrast to that
of the leopard ( Panthera pardus ), side-striped jackal
( Canis adustus ), and serval cat ( Felis serval ), which
showed average persistence. As in other studies ( e.g.,
Struhsaker & Oates 1995; Bodmer et al. 1997 ), I found
that herbivores including colobid monkeys were also
among those most prone to extinction.
Species most preferred by hunters and consumers
were as likely as less-preferred species to become locally
extinct (Fig. 4). Other studies in the tropics also suggest
Correlates of Extinction in West Africa 741
that hunters are unselective of prey (Bodmer 1995; Fa et
al. 1995; FitzGibbon 1998 ) but maximize their harvest
by extracting any animal of value when they encounter
it (Alvard 1998; J. Brashares, unpublished data). My esti-
mates of human preference were based on the eco-
nomic and subsistence value of prey as determined by
surveys of markets and hunters, but these estimates do
not account for the degree to which species are targeted
as a result of the threat they pose as crop raiders or pred-
ators of livestock and people. Combining data on the
economic value of prey and human-wildlife conflicts
may produce a better predictor of species persistence.
Species’ Persistence in Ghana and Broader Trends
Species persistence rates in Ghana and IUCN Red Data
Book listings of species global vulnerabilities were
highly correlated ( Fig. 5 ). Assuming that Ghana com-
prises only a small part of the ranges of species used in
such a comparison (Ghana is 3% of the range of 40 of
the 41 species considered), the positive correlation be-
tween the global status of species and their persistence
in Ghana suggests that the traits that predicted species
vulnerability in Ghana are also useful predictors else-
where.
I have drawn out common results from studies of ex-
tinction in undisturbed habitat islands and those from
studies of the persistence of commercially valuable spe-
cies under exploitation. The difference between these
approaches is that the first attempts to comprehend
how ecological and life-history traits affect the extinc-
tion of species in the absence of modern humans. By
studying extinction in exploited systems, however, we
may also learn how exploitation by humans interacts
with ecological, behavioral, and life-history traits to af-
fect persistence. In practice, separating “natural” extinc-
tions from anthropogenic ones will be difficult for many
reasons, including the fact that exploitation by humans
often acts on the same traits of species as natural pro-
cesses. For example, body size, trophic level, and rarity
have been associated with the vulnerability of species to
natural extinction but might also predict extinction in
hunted habitats for purely economic reasons. Neverthe-
less, studies of natural extinction have typically ap-
peared in journals favoring topics in theoretical evolu-
tionary ecology, whereas studies of extinction in
exploited systems have appeared more often in applied
ecology and conservation biology journals. As evinced
here, these two types of study may provide similar rec-
ommendations for managers of species and habitats.
Acknowledgments
The Ghana Wildlife Division graciously permitted me to
work in Ghana’s national parks and reserves. In particu-
Conservation Biology
Volume 17, No. 3, June 2003
742 Correlates of Extinction in West Africa
lar, I thank M. K. Sam, N. Ankudey, and B. Volta for ac-
cess to historical data, research permits, letters of intro-
duction, and advice. I thank C. Kresge, P. Kresge, J.
Mason, and the staff of the Nature Conservation Re-
search Centre for logistical support in the field. I thank
P. Arcese for helpful comments throughout the course
of this study; P. B. Coppolillo, M. Kilpatrick, K. Martin,
B. Sandercock, A. R. E. Sinclair, J. N. M. Smith, and two
anonymous reviewers for helpful comments on the
manuscript; and T. Garland, A. Kozak, and V. LeMay for
advice on statistical methodology. This work was sup-
ported by the Department of Wildlife Ecology at the Uni-
versity of Wisconsin, the Centre for Applied Conserva-
tion Biology at the University of British Columbia, the
Zoological Society of Milwaukee County, National Sci-
ence Foundation grant IBN-9458122 to P. Arcese, and
the Killam Trusts.
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