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Animal Conservation. Print ISSN 1367-9430

Will their armaments be their downfall? Large horn size

increases extinction risk in bovids
J. Bro-Jørgensen
Mammalian Behaviour & Evolution Group, Department of Evolution, Ecology & Behaviour, Institute of Integrative Biology, University of
Liverpool, Leahurst Campus, Neston, UK

Keywords
extinction risk; ungulates; conservation
management; sexual selection; sexual size
dimorphism; selective harvesting;
overexploitation; climate change.
Correspondence
Jakob Bro-Jørgensen, Mammalian
Behaviour & Evolution Group, Department
of Evolution, Ecology & Behaviour, Institute
of Integrative Biology, University of
Liverpool, Leahurst Campus, Neston
CH64 7TE, UK. Tel: +44 (0) 151 794 6009;
Fax: +44 (0) 151 794 6107. Email:
bro@liv.ac.uk
Editor: Iain Gordon
Associate Editor: Hervé Fritz
Received 12 June 2012; accepted 15 May
2013
doi:10.1111/acv.12062
Abstract
Clarifying the emergent fitness associated with sexually selected traits under the
current, increasingly anthropogenic selection regimes is important to understand
ongoing evolutionary changes in nature and inform the conservation management
of endangered species. Several reasons exist why sexual selection may affect
extinction risk. Increased risk may result either from inherent trade-offs between
sexually selected traits and viability traits or from selective hunting of sexually
selected species. Reduced risk is also possible, for instance if the preference for
high-performing mates characteristic of sexually selected species has beneficial
genetic consequences for the population. Here, I show that the threat level of
bovid species increases with large male horn size. This is the first time, to my
knowledge, that sexually selected weaponry has been shown to increase extinction
risk at the interspecific level. However, threat level was unrelated to another trait
under sexual selection, sexual body size dimorphism, indicating that the effect of
sexual selection on extinction risk depends on trait-specific interactions with
extrinsic factors. The results suggest that the higher threat level of long-horned
species is not primarily due to current trophy hunting practices and rather point to
environmentally induced viability costs as a possible main driver. Still, the fact
that long-horned species are known to be preferred by trophy hunters highlights
the importance of continuously monitoring trophy hunting practices to assure
their long-term sustainability.

Introduction
Selection favours individuals who maximize their fitness by
striking an optimal balance between investment in repro-
duction and survival (Stearns, 1992). However, awareness is
increasing that selection operates at multiple levels, and
traits evolved by individual selection in relatively stable
natural environments may become disfavoured by selection
at the species level when selection regimes change
(Jablonski, 2008; Rabosky & McCune, 2010). In the
present, increasingly human-dominated environment, iden-
tification of traits associated with elevated extinction risk
can therefore provide important information on current
threat processes and help guide priorities for conservation
management. Particular attention has centred on the poten-
tial impact that sexually selected traits may have on popu-
lation viability (Morrow & Pitcher, 2003; Candolin &
Heuschele, 2008). A classical, but largely anecdotal example
is provided by the Irish elk, Megaloceros giganteus, whose
palmate antlers, spanning up to 3.6 m, are believed to have
evolved primarily through intrasexual selection (Gould,
1974; Barnosky, 1985). Symptomatic of the wider debate,
controversy has arisen over the role of the antlers in the
extinction of the deer around 7500 BP, at a time when its
habitat was shrinking and hunting intensifying following the
last glacial (Moen, Pastor & Cohen, 1999; Gonzalez,
Kitchener & Lister, 2000; Stuart et al., 2004; O’Driscoll
Worman & Kimbrell, 2008). Speculations on the
maladaptiveness of such extreme cranial appendages may
seem intuitively appealing considering the likely costs
involved; however, robust evidence that sexually selected
weaponry can indeed increase extinction risk is lacking.
Theoretically, sexual selection can both enhance and
reduce extinction risk. Higher extinction risk may be pre-
dicted where environmental changes alter the trade-offs
between natural selection and sexual selection: when the
environment deteriorates, increasing viability costs can
render sexually selected traits maladaptive (Kokko &
Rankin, 2006). For example, although several intraspecific
studies on male bovids have linked large horn size to
increased individual survival in stable environments
(Bergeron et al., 2008; Bonenfant et al., 2009), a study on
the Soay sheep, Ovis aries, has revealed that the fitness of
long-horned rams is above average only under benign

80 Animal Conservation 17 (2014) 80–87 © 2013 The Zoological Society of London

J. Bro-Jørgensen
environmental conditions: under harsh conditions, reaction
norms cross and long-horned rams perform below average
(Robinson et al., 2008). At the interspecific level, such envi-
ronmentally induced costs may translate into higher extinc-
tion risk of long-horned species under ecological stress as
even relatively short-horned individuals of long-horned
species may be more affected than the most long-horned
individuals of short-horned species.
Higher extinction risk of sexually selected species may
also be consequential to human exploitation when hunting
directly targets specific sexually selected traits, such as
manes (Whitman et al., 2004), tusks (Milner-Gulland &
Mace, 1991), elaborate plumage (Metz & Ankney, 1991) or
indeed horns (Johnson et al., 2010). Often, however, such
selective harvesting is biased towards males, and since
females are typically the sex limiting reproductive rates
(Emlen & Oring, 1977), the impact on population viability
may be ameliorated (Caro et al., 2009). Still, in extreme
cases, both the number and quality of males in the mating
pool can drop below the critical level at which recruitment is
depressed, potentially leading to reproductive collapse
(Milner, Nilsen & Andreassen, 2007). The impact of male
removal may be compounded where hunters specifically
target the reproductively most important individuals by
selecting those with the most extreme expression of the sexu-
ally selected trait (Annighoefer & Schuetz, 2011).
The very process of sexual selection may as well be intrin-
sically linked to reduced population viability because the
associated restrictive mating patterns both promote
inbreeding and inflict a selection load which slows down the
rate of adaptation (Kokko & Brooks, 2003; Candolin &
Heuschele, 2008). Alternatively, however, it is also possible
that selective mating characteristic of sexually selected
species has a positive, rather than a negative, impact on
population viability: the widespread preference for high per-
forming mates has the potential to purge populations of
deleterious alleles and accelerate the rate of adaptation
to environmental changes (Candolin & Heschele, 2008;
Jarzebowska & Radwan, 2010).
Given this diversity in theoretical possibilities, there is a
need to establish how sexually selected traits actually relate
to extinction risk under contemporary selection regimes,
and here the family Bovidae presents an exceptional oppor-
tunity. This speciose family is an urgent conservation prior-
ity with 40% of the extant species threatened with extinction,
the main threats being habitat loss and exploitation for
diverse products such as meat, horns, hides and wool (http://
www.iucnredlist.org on 15 October 2012). Bovid species
vary dramatically in their expression of sexually selected
traits. Thus, the horns of males as well as the often conspicu-
ous exaggeration of male body size (reflected in sexual body
size dimorphism, SSD) are generally believed to have
evolved under the influence of sexual selection due to their
function in intrasexual competition for mates (Lundrigan,
1996; Loison et al., 1999; Pérez-Barbería, Gordon & Pagel,
2002; Caro et al., 2003; Bro-Jørgensen, 2007). Both these
traits can incur significant survival costs (Owen-Smith,
1993; Andersson, 1994): large male body size, which
Animal Conservation 17 (2014) 80–87 © 2013 The Zoological Society of London
Large horn size increases extinction risk in bovids
increases energy demands, can lead to foraging strategies
that enhance mortality (Hay, Cross & Funston, 2008), and
large male horns may not only impose physical and physi-
ological constraints (Geist, 1966; Picard, Festa-Bianchet &
Thomas, 1996), but also be directly targeted by hunters
(Johnson et al., 2010; Palazy et al., 2012).
Using the comparative method (Harvey & Pagel, 1991;
Fisher & Owens, 2004), I here investigate the effect of sexual
selection on extinction probability among bovid species by
testing how threat level responds to the expression of the
two sexually selected traits, male horn length and SSD. If
sexual selection in general increases extinction risk due to
trade-offs with viability selection, I predict that threat level
will increase with both male horn length and SSD. Con-
versely, if sexual selection in general reduces extinction risk
by accelerating adaptive change, I predict that threat level
will decrease with both these traits. If the two traits differ in
their effect on extinction risk due to trait-specific interac-
tions with extrinsic factors, I predict that they will relate
differently to threat level. Specifically, if an association
between extinction risk and large male horn size is due to
current trophy hunting practices, I predict that (1) trophy
species will possess longer horns, (2) long-horned species
will be more exposed to overexploitation, (3) trophy species
will be more threatened, and (4) trophy value will increase
with horn length. Alternatively, if a link between extinction
risk and horn size is an indirect consequence of long-horned
species occupying more open habitats where they may be
easier to hunt and more exposed to threats such as habitat
fragmentation, I predict that (1) long-horned species should
predominate in open habitats, and that (2) species in open
habitats should be more threatened. Finally, as large-
bodied, slow reproducing species are often at higher extinc-
tion risk (Cardillo et al., 2005), I also test whether large
body size increases extinction risk.
Materials and methods
Data
The data collection focused on all extant bovid species as
well as those which have gone extinct within the current
extinction crisis (< 400 years). Information was gathered on
the following morphological and ecological variables:
average (avg.) male body mass, avg. female body mass, avg.
male horn length, avg. male shoulder height and habitat
density. Information was available for 119 out of 140
species, including two out of four extinctions, that is the
auroch Bos primigenius (†1627) and the bluebuck
Hippotragus leucophaeus (†1800, approx.). Species-specific
body mass was calculated as the mean of male and female
body masses, and sexual body size dimorphism was calcu-
lated as the ratio of male to female body mass. Relative male
horn length was calculated by dividing horn length, a linear
measure, by the most readily available linear measure of
body size, that is, shoulder height (Jarman, 1983). All mor-
phological measures were log-transformed prior to analysis
to make them suitable for regression analysis. Following
81
Large horn size increases extinction risk in bovids
Bro-Jørgensen (2008), habitat density was coded as a binary
categorical variable where open habitats (0) included
deserts, steppes, mountains, all types of savannas and open
woodlands, and closed habitats (1) included thickets, dense
woodland and forests. The data on the morphological meas-
urements and habitat was copied from Bro-Jørgensen (2007,
2008) and supplemented with information on (1) shoulder
height and (2) additional species from the sources listed in
the online Supporting Information.
Extinction risk was coded as a continuous variable based
on threat status on the International Union for Conserva-
tion of Nature (IUCN) red list (accessed 9 November 2011
on http://www.iucnredlist.org) as follows: 1 = ‘least con-
cern’, 2 = ‘near threatened’, 3 = ‘vulnerable’, 4 = ‘endan-
gered’, 5 = ‘critically endangered’ and 6 = ‘extinct’ or
‘extinct in the wild’ (Fisher & Owens, 2004). Whether a
species was listed as threatened by overexploitation on the
red list was coded as a binary categorical variable. For
overexploited species, the IUCN classification does not
systematically give information on the exploited animal
product, and I therefore categorized species currently
subjected to trophy hunting (henceforth ‘current trophy
species’) based on the comprehensive hunting company
survey of Palazy et al. (2012). Trophy value (US$) was also
obtained from Palazy et al. (2012); where several subspecies
were listed, the price was averaged.
Comparative analysis
All analyses applied the interspecific comparative method,
including only evolutionarily independent contrasts
between species (Harvey & Pagel, 1991; Fisher & Owens,
2004). The comparative analyses were performed in R (R
Development Core Team, 2011) using the packages caper
(Orme et al., 2011) and ape (Paradis, Claude & Strimmer,
2004). Phylogenetic relatedness was controlled for by
assuming a gradual model of evolution based on the
phylogenetic supertree and branch lengths in Fernández &
Vrba (2005). Contrasts at polytomies were calculated fol-
lowing the method described by Pagel (1992). Independent
contrasts were calculated from the crunch algorithm where
all traits were continuous, and the brunch algorithm where
a categorical trait was included (Orme et al., 2011);
phylogenetic generalized least squares analysis gave similar
results. All regressions were done through the origin, with
Table 1 Phylogenetic correlations between extinction risk in bovids and the test variables
Independent Multivariate analysis
variables Coefficient (B) d.f. t P
Relative male 2.153 (1.737) 1,93 (1,90) 2.32 (2.30)
horn lengtha
Sexual body size −0.566 (−0.662) 2,92 (2,89) −0.58 (−0.84)
dimorphism
Body mass −0.098 (0.042) 3,91 (3,88) −0.38 (0.20)
a
Horn length/shoulder height.
Numbers in parenthesis refer to the results obtained after removal of outliers, see ‘Materials and methods’.
82
J. Bro-Jørgensen
backward elimination of non-significant variables in multi-
ple regressions.
Crunch standardizes contrasts between taxa by dividing
them by their expected standard deviation, estimated as the
square root of the sum of the branch lengths between each
taxon and their common ancestor. The approach assumes
that the evolution of continuous traits can be modelled as a
random walk process (Felsenstein, 1985), and this assump-
tion was tested by regressing the estimated contrasts against
their estimated nodal values (Orme et al., 2011). I also tested
the success of the contrast standardization procedure by
regressing the absolute values of the standardized contrasts
against the estimated standard deviation at each node,
thereby assessing the heterogeneity of variance in the residu-
als and the suitability of the data for regression analysis
(Orme et al., 2011). All of these tests were non-significant as
expected given the assumptions, except in one case where
SSD was included as the independent variable in a bivariate
regression. Following Møller (2011), I therefore checked the
robustness of the results by (1) repeating the analyses with
the independent variables expressed as ranks, and (2) using
the caic.robust command to remove outliers with absolute
studentized residuals exceeding three (Jones & Purvis, 1997).
Neither of these alternative approaches changed the results
qualitatively.
Due to correlations between independent variables, I
assessed the potential effect of collinearity on the results of
the multivariate analyses by calculating tolerance and vari-
ance inflation factors using the Predictive Analytics
SoftWare Statistics 17.0.2. In all cases, the variance inflation
factors were < 1.25 and thus well below the commonly
accepted critical thresholds for significant collinearity of
5–10 (McClave & Sincich, 2003).
Results
Relative male horn length was found to be a significant pre-
dictor of current extinction risk in bovids (Table 1; Fig. 1a).
By contrast, sexual body size dimorphism did not show any
demonstrable relationship with extinction risk, whether ana-
lysed in bivariate or multivariate regression models (Table 1;
Fig. 1b). This suggests that increased extinction risk is not a
general consequence of sexual selection.
Current trophy species were not characterized by particu-
larly long male horns (whether a species was listed as a
Bivariate analyses
Coefficient (B) d.f. t P
0.022 (0.024) (As multivariate analysis)
0.563 (0.404) −0.873 (−0.791) 1,93 (1,91) −0.96 (−1.02) 0.340 (0.310)
0.705 (0.842) 0.048 (0.129) 1,93 (1,91) 0.22 (0.71) 0.823 (0.482)
Animal Conservation 17 (2014) 80–87 © 2013 The Zoological Society of London
J. Bro-Jørgensen Large horn size increases extinction risk in bovids

a trophy species as a function of relative male horn length:

2.0 d.f. 1,27, t = 0.63, P = 0.534), nor could any link between
large male horn size and exposure to overexploitation be
demonstrated (whether a species was listed as threatened by
Extinction risk

1.0
overexploitation by IUCN as a function of relative male
horn length: d.f. 1,18, t = 0.25, P = 0.808). Thus, in spite of
0.0 including 6 of 20 endangered, and 8 of 20 vulnerable species,
current trophy species were generally less, rather than more,
threatened than other species (threat level as a function of
-1.0 whether a species was listed as a trophy species: d.f. 1,27,
t = −5.24, P < 0.001). Male horn length was, however, posi-
tively linked to trophy value (trophy value as a function of
-2.0
0.00 0.05 0.10 0.15 0.20 relative male horn length: d.f. 1,61, t = 2.09, P = 0.041), con-
Relative male horn length sistent with an earlier reported preference for species with
large horn size among trophy hunters (Palazy et al., 2012).
Overall, these findings speak against trophy hunting, as
b reflected in current practices, as the main driver of the
2.0 increased threat levels of long-horned bovids.
Although long-horned species showed a weak, non-
significant tendency to inhabit more open environments
Extinction risk

1.0 (relative male horn length as a function of whether a species

occurred in open or closed habitat: d.f. 1,11, t = −1.70,
P = 0.117), open habitat species tended to be less, rather
0.0
than more, threatened than closed habitat species (threat
level as a function of whether a species occurred in open or
-1.0 closed habitat: d.f. 1,11, t = 2.10, P = 0.060). This last result
contradicts that increased habitat fragmentation, or facili-
tated hunting, in open habitats should indirectly drive the
-2.0 higher extinction risk of long-horned species. By default, the
-0.25 -0.15 -0.05 0.05 0.15 results are most consistent with environmentally induced
Sexual size dimorphism viability trade-offs of large horn size as the main driver of
the relationship between extinction risk and horn length.
No association was found between body size and the
c extinction risk of bovids (Table 1; Fig. 1c).
2.0

1.0
Extinction risk

Discussion
The link between the horn length of male bovids and
0.0 increased extinction risk is a novel discovery suggesting that
sexually selected weaponry can adversely affect species sur-
vival. However, the lack of relationship between sexual
-1.0 body size dimorphism and extinction risk suggests that the
intensity of sexual selection per se is not a reliable predictor
of extinction probability. This conclusion is consistent with
-2.0 previous studies on artiodactyls. Morrow & Fricke (2004)
-0.6 -0.3 0 0.3 0.6
found no association between extinction probability and
Body mass either of the two sexually selected traits (testis size and
SSD), and among Ghanaian ungulate populations,
Figure 1 Predictors of extinction risk in bovids. (a) Contrasts in extinc-
Brashares (2003) found a negative rather than a positive link
tion risk against contrasts in relative male horn length (i.e. horn
between the vulnerability to local extinction and polygyny, a
length/shoulder height). The line shows the significant slope from
key determinant of sexual selection (Pérez-Barbería et al.,
linear regression through the origin; the slope remained significant
after removal of the three outliers denoted by open squares (Table 1).
2002; Shuster & Wade, 2003). The costs and benefits of
(b) Contrasts in extinction risk against contrasts in sexual size dimor- sexually selected traits thus appear to show a significant
phism. (c) Contrasts in extinction risk against body mass contrasts. In degree of trait-specificity; in particular, it is possible that
(b) and (c), no significant slopes were obtained from linear regression SSD provides context-dependent benefits when resources
through the origin. are scarce by allowing the sexes to exploit distinct, size-
dependent ecological niches (Ruckstuhl & Neuhaus, 2000).

Animal Conservation 17 (2014) 80–87 © 2013 The Zoological Society of London 83

Large horn size increases extinction risk in bovids
Why then is extinction risk specifically linked to male
horn size? No evidence was found that bovids currently
listed as trophy species by hunting companies are more
exposed to overexploitation at present. This is in agreement
with some recent case studies reporting only limited evi-
dence of unsustainability in bovid trophy quotas (e.g. Caro
et al., 2009; Atickem et al., 2011). However, such case
studies should not be freely generalized across taxa or
regions (Milner et al., 2007; e.g. Annighoefer & Schuetz,
2011), and in the present study I was unable to take into
account illegal trophy hunting of threatened species on
which data are hard to come by. To facilitate future evalu-
ation of the conservation impact of trophy hunting, I there-
fore recommend that the IUCN commence to systematically
categorize overexploited species according to the nature of
the product targeted, in the case of bovids including (1)
horns, and whether these are used (a) as trophies or (b) in
traditional medicine, (2) meat, (3) hides and (4) wool.
Moreover, it cannot be ruled out based on the present study
that less conservation-friendly trophy hunting practices in
the past could have contributed to the higher extinction risk
associated with large male horn size today.
Still, the absence of current evidence that over-
exploitation links horn size to extinction risk increases the
focus on habitat-related factors as the primary cause. No
support was found for the specific idea that threats associ-
ated with open habitats indirectly link long-horned species
to higher extinction risk, and the available evidence thus
leaves as the presently most parsimonious explanation that
large horn size primarily promotes extinction because of
viability trade-offs that intensify under environmental stress
[Fig. 2 (i–ii)]. The costliness of large horns in adverse envi-
ronments can be explained by a negative impact on fitness
components such as the ability to acquire adequate
resources or escape predators (Geist, 1966; Robinson et al.,
2008). The higher extinction risk of long-horned species is
possibly a natural phenomenon due to the dynamic nature
of the wild ecosystems inhabited by bovids; yet, the largely
human-caused degradation of bovid habitats at present
could affect long-horned species disproportionately. The
simultaneous importance of climatic and anthropogenic
drivers has indeed been highlighted by studies of late qua-
ternary extinctions (Stuart et al., 2004; Prescott et al.,
2012a), where more recent examples include several long-
horned species, such as the long-horned buffalo, Pelorovis
antiquus and the giant hartebeest, Megalotragus priscus on
the African savannah (†11 000 BP; Klein, 2000; Gentry,
2011), and the giant bison, Bison latifrons in North America
(†20 000 BP; Scott, 2010).
Even if no causal link between trophy hunting and the
higher extinction risk of long-horned species is suggested by
the present findings, the documented preference for long-
horned species among trophy hunters (Johnson et al., 2010;
Palazy et al., 2012) still underscores the importance of
rigorously monitoring trophy hunting practices in terms
of their long-term sustainability (Bunnefeld, Hoshino &
Milner-Gulland, 2011; see also Harris & Pletscher, 2002;
Damm, 2008; Sachedina & Nelson, 2010). Where hunting is
84
J. Bro-Jørgensen
Horn size
+ (iii)
+ (i) Hunter
preference
(iv)
+
Increased
suscepƟbility to + (vi) Intensifying
habitat decline exploitaƟon
(ii) +
+ (v)
Rarity
+ (vii)
ExƟncƟon risk
Figure 2 Proposed links between trophy size and extinction risk.
Higher susceptibility to habitat deterioration in species with large
horn size (Robinson et al., 2008) (i) can lead to population declines
and hence promote rarity (ii). Hunter preference for large trophy size
(Johnson et al., 2010; Palazy et al., 2012; this study) (iii) also makes
these species particularly likely to be targeted by exploitation (iv); if
poorly regulated, trophy hunting can therefore drive rarity as well (v).
Because hunters also prefer rare species (Johnson et al., 2010;
Palazy et al., 2012) (vi), positive feedback between rarity and hunter
preference can moreover result in ‘vicious circles’ of intensifying
exploitation (Palazy et al., 2012) (iv,v,vi). As a consequence of these
forces, long-horned species are at elevated risk of extinction, as
demonstrated in this study (vii).
poorly regulated, this preference may otherwise cause popu-
lation declines in the most vulnerable species [Fig. 2 (iii–iv)].
An independent preference for rare species, especially
among experienced hunters (Lindsey et al., 2006; Johnson
et al., 2010; Prescott et al., 2012b), has the potential to accel-
erate population declines by generating a positive feedback
cycle between hunter preference and rarity, thereby leading
to an anthropogenic Allee effect [Courchamp et al., 2006;
Palazy et al., 2012; Fig. 2 (iv–vii)]. Moreover, poorly
managed trophy hunting can have adverse consequences by
promoting unnatural directional selection for shorter horn
length (Coltman et al., 2003; Mysterud, 2011). On the other
hand, when properly managed, trophy hunting can bring
significant conservation benefits by translating ecosystem
services into economic benefits for local communities (Lewis
& Alpert, 1997; Loveridge, Reynolds & Milner-Gulland,
2007; Frost & Bond, 2008; Naidoo et al., 2011), especially
where alternative land use strategies, such as ecotourism,
cannot generate sufficient income to ensure preservation
(Wilkie & Carpenter, 1999; Lindsey et al., 2006).
Animal Conservation 17 (2014) 80–87 © 2013 The Zoological Society of London
J. Bro-Jørgensen
Long-horned species may also be particularly vulnerable
to unsustainable use of horns from wild bovids in tradi-
tional medicine (e.g. in kouprey Bos sauveli, markhor Capra
falconeri, mountain nyala Tragelaphus buxtoni and saiga
Saiga tatarica; http://www.iucnredlist.org on 15 October
2012). Without evidence that such exploitation can be
managed for a net conservation benefit, and in the absence
of any medical justification, a strong case exists for discour-
aging this practice in favour of more effective and
conservation-friendly medical treatments.
No association was found between body size and extinc-
tion risk in bovids. This may seem surprising since large-
bodied mammals, with their low reproductive rates, are
often more extinction prone (Cardillo et al., 2005).
However, the negative finding mirrors previous studies of
ungulates, thus confirming that predictors of threat in
mammals show a high degree of taxon specificity (Price &
Gittleman, 2007; Cardillo et al., 2008). The lack of relation-
ship may be related to the tendency towards higher threat
levels of bovids in closed habitats, where species are gener-
ally smaller (Bro-Jørgensen, 2008). In tropical forests,
overexploitation of bovids for bushmeat is a main concern
(Fa & Brown, 2009), and unsustainable subsistence hunting
of smaller species may cancel out the generally positive
effect of small body size on ecological resilience.
In conclusion, the present study shows that sexually
selected weaponry can increase extinction risk but also that
the effect of sexual selection on extinction risk is trait spe-
cific. Pinpointing the particular circumstances under which
sexually selected traits are associated with increased extinc-
tion risk is therefore important for the conservation
management of endangered species. The discovery that
extinction risk in bovids increases with male horn size
underscores the importance of monitoring trophy hunting
due to a preference among trophy hunters for long-horned
species. Although the findings suggest that the higher threat
level of long-horned species is not primarily driven by
current trophy hunting practices, further studies are needed
on mortality causes of wild bovids in order to establish more
firmly the relative importance of natural and anthropogenic
threat drivers.
Acknowledgements
I am grateful to Mogens Andersen and Hans J. Baagøe at
the Zoological Museum of Copenhagen for their excep-
tional assistance. I thank the Mammalian Behaviour and
Evolution Group at Liverpool University, Tim Caro and
two anonymous reviewers for valuable comments. This
research was supported by an RCUK fellowship.
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Supporting information
Additional Supporting Information may be found in the
online version of this article at the publisher’s web-site:
Appendix S1. Additional data sources.
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