2019

What are the evolutionary drivers that

have resulted in the development of
horns in female bovids?
PREDATION INTERACTIONS, NATURAL SELECTION AND EVOLUTION
HAWKES, SARAH, (MISS) (S217174019)
Table of Contents
Abstract ............................................................................................................................... 2

Introduction ......................................................................................................................... 2

Male Vs Female Horns ........................................................................................................ 3

Male Horns....................................................................................................................... 3

Female Horns .................................................................................................................. 3

Hypotheses ......................................................................................................................... 3

Male Mollification Hypothesis ........................................................................................ 3

Competition Hypothesis ................................................................................................. 3

Anti-Predator Defence Hypothesis ................................................................................ 4

Territoriality Hypothesis ................................................................................................. 4

Intrasexual Selection Hypothesis .................................................................................. 4

Female horns in bovids ...................................................................................................... 4

Traits and Drivers............................................................................................................ 4

Territoriality ................................................................................................................... 4
Body Size and Conspicuousness .................................................................................. 5

Mechanism of Trait Evolution ........................................................................................ 5

Examples ............................................................................................................................. 5

Prey morphology drive predator prey preferences in Acinonyx jubatus .................... 5

Discussion and Conclusion ............................................................................................... 6

Acknowledgements ............................................................................................................ 7

References .......................................................................................................................... 7

1
What are the evolutionary drivers that have
resulted in the development of horns in
female bovids?
Sarah Hawkes
217174019

Abstract
Weaponry such as horns is present in majority of male bovids which is driven by intrasexual
selection, but the seemingly random appearance of horns in some female bovids is still
unknown. This paper aimed to review the driving forces behind the evolution of horns in some
female bovids but not in others. A variety of hypotheses have been proposed to account for
these inconsistencies. Such hypotheses include male mollification, competition for food,
territoriality and defence against predators. The main driving force behind the evolution of
horns in female bovids is natural selection, which selects for the beneficial traits of horns such
as protection, in order to improve the fitness of the individual female so that they would survive
and reproduce. It was concluded that anti-predatory defence is the most likely hypothesis
explaining the presence of horns in most female bovids.

Introduction
Predation interactions result in an evolutionary response in the prey species, through natural
selection, which selects upon traits that improve the reproductive and survival success of an
individual (Abrams, 1990). Most animals are preyed upon by predators. when a prey
population is exposed to a predator, evolutionary responses can occur as a result of natural
selection, to improve the fitness of the prey population to defend themselves against the
predator (Abrams, 1990). Majority of male bovids display horns as a result of intrasexual
selection, but in females only a few exhibits the presence of horns. These inconsistencies
have resulted in a variety of hypotheses being formulated to explain the presence of horns in
some female bovids and not others, as well as the different functions male and female horns
provide. This paper aimed to review the driving forces behind the evolution of horns in some
female bovids and not others, as well as studying the relationship between horns and predator-
prey interactions.

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Male Vs Female Horns
Male Horns
The horns present in male and female bovids are shaped differently from one another and
possess different functions (Packer, 1983). Majority of male bovids exhibit the presence of
horns as a result of intrasexual competition. Male bovids have different shaped horns
dependent upon the style of fighting they make use of, for example, if the bovid exhibits
behavior such as ramming of the opponent, their horns will have tips that face downwards
(Caro, et al., 2003).

Female Horns
The horns present on female bovids are often smaller in size and shaped differently to the
horns present on males (Stankowich & Caro, 2009). Hypotheses for this is male mollification,
competition for food, territoriality and defence against predators (Stankowich & Caro, 2009).
The horns found in female bovids tend to be twisted on the large species in order to wrestle
other individuals or they are shaped straight and face backwards in order to stab predators
(Caro, et al., 2003).

Hypotheses
Male Mollification Hypothesis
The presence of horns in some female bovids is a form of mimicry of the male bovids in order
to appease the male’s aggression towards their male offspring (Stankowich & Caro, 2009).
This, therefore, prolongs the amount of time their male offspring remain within the herd as well
as ensures the reproductive success and survival of the male offspring (Estes, 1991). There
is a general trend whereby the female bovids that are part of a mixed-sex group will be more
likely to exhibit horns than female bovids that are in a single sex groups (Stankowich & Caro,
2009).

Competition Hypothesis
This hypothesis suggests that the presence of horns in females could have evolved as a result
of intrasexual competition for resources (Roberts, 1996). The amount of competition that is
displayed is directly correlated with the size of the group (Roberts, 1996). It therefore, is
suggested that the larger the group size of females the higher the probability that they will
possess horns (Roberts, 1996). It can be further deduced that females compete for resources
in order to ensure that they obtain maximum reproductive benefits (Stockley & Bro-Jørgensen,
2011).

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Anti-Predator Defence Hypothesis
The presence of horns in some females and not others may be as a result of the horns
functioning as a defence against predation (Kodric-Brown & Brown, 1984). Larger females will
benefit more from the possession of horns than smaller females as a result of the energy
requirements needed to develop horns (Stankowich & Caro, 2009). Female horns face
backwards and are shaped straight, which suggest that they are specifically adapted for
piercing predators (Packer, 1983). Openness of habitat also plays a role in anti-predatory
defence, as animals that live in areas where they are large and exposed will not be able to
find refuge to avoid predators, therefore, they require another mechanism of protection.
Through this lack of refuge horns have evolved in order to provide protection from predators
(Stankowich & Caro, 2009).

Territoriality Hypothesis
It is suggested that females that compete with each other over territory would receive a greater
benefit from the presence of horns than those females that did not compete over territory
(Stankowich & Caro, 2009). The presence of horns in territorial females, would improve their
ability to defend their territory as they will have an advantage over the females that do not
possess horns. Intrasexual competition over territories resulted in the development of horns.

Intrasexual Selection Hypothesis

Majority of male bovids exhibit the presence of horns as a result of intrasexual selection
(Stankowich & Caro, 2009). In males, the horns are shaped so that they may ward off an
attack as well as fight an opponent, which suggests that they are mainly present for intrasexual
combat (Stankowich & Caro, 2009). Intrasexual combat in males takes place in order to attract
a mate, horns improve the males chances of winning the fight, which in turn improves his
chances of finding a mate and passing on his genes (Kappeler, 1996).

Female horns in bovids

Traits and Drivers
Territoriality
Females that compete with other females and display territoriality tend to exhibit the presence
of horns, whereas females that are not territorial do not (Stankowich & Caro, 2009). It can
therefore, be deduced that territoriality acts as a driver for the evolution of horns in females.
The horns were selected for in order to improve their ability to protect their territory.

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Body Size and Conspicuousness
Larger sized female bovids are characteristically better at defending themselves against
predators than smaller sized female bovids (Stankowich & Caro, 2009). It was found that
intermediate sized prey displays the presence of horns, suggesting, that they are just under
the size threshold whereby predators avoid prey; therefore, they require another mechanism
of protection which comes in the form of horns. Smaller sized animals are able to flee quickly
and take refuge, thus the energy requirements necessary for the development of horns
outweigh the need for horns. Body size therefore, acts as a driver of horn evolution. There is
one exception to this driver that is found in Tibetan antelope whereby, they are large in size
and live in areas where they are exposed and yet lack the presence of horns, this is due to
the fact that they dig holes in the desert and then lay down in them, thus, reducing their
exposure to predators (Stankowich & Caro, 2009).

Mechanism of Trait Evolution

Natural selection is the main driving force behind the evolution of horns in female bovids. The
allele frequencies within the population will change over a relatively short period of time as a
result of natural selection. Natural selection will act by either selecting genes that are pre-
existing or it will select newly arisen mutations (Barrett & Schluter , 2008). The process
involves individuals that have specific beneficial traits within a population that exhibit a higher
reproductive and survival success than other individuals, and, as a result of this pass these
genes onto their offspring (Hancock, et al., 2010). The presence of horns in the population of
females is considered a beneficial trait that is selected on and passed onto their offspring as
they are better adapted at defending themselves from predators, therefore, they will exhibit a
higher fitness.

Examples
Prey morphology drive predator prey preferences in Acinonyx jubatus
A study was conducted on the Cheetah (Acinonyx jubatus) in order to analyze the predator-
prey interaction. The study showed that the presence of horns in the prey influences the
cheetahs prey preference. It was found that horns in females were present in the medium
sized individuals that were short of being too big for the cheetah to prey on (Clements, et al.,
2016).

A predator will choose its prey based on the energetics cost of catching it, as well as the
probability of injury. The risk of predation is a function of the interaction between the size of
the prey and the presence of weaponry (Stankowich, 2011). The morphological traits of prey,

5
predator behavior and density of predators have a large influence on the predator’s prey
preferences. It was found that upper prey thresholds were identified and that larger body sizes
serve to provide protection to the prey species (Stankowich, 2011). The study provides
definitive evidence that horns are a successful antipredator mechanism (Clements, et al.,
2016).

Discussion and Conclusion

This paper aimed at determining the driving forces behind the evolution of horns in some
female bovids and not others as well as the relationship between horns and predator-prey
interactions. A variety of hypotheses have been proposed to account for these observations.
Such hypotheses include male mollification, competition for food, territoriality and defence
against predators.

Natural selection is the main driving force behind the evolution of horns in female bovids
(Barrett & Schluter , 2008). Through natural selection the trait for horns was selected as a
result of predators, territoriality, body size and the individual’s ability to find refuge (Stankowich
& Caro, 2009). The trait was selected for in order to improve the fitness of the individual so
that they are better adapted to survive and reproduce (Barrett & Schluter , 2008). These
advantageous traits will then be passed on to their offspring resulting in the population survival
rates improving (Barrett & Schluter , 2008).

The hypothesis that best explains the presence of horns in some female bovids and not others
is the anti-predator defence hypothesis (Kodric-Brown & Brown, 1984). This hypothesis is
driven by body size, conspicuousness and openness of habitat, therefore, incorporating a
variety of drivers into one hypothesis. It was supported by results obtained by Stankowich &
Caro (2009) in which body size, openness of habitat and conspicuousness had the highest
probability for the presence of horns in female bovids. Male mollification, competition for
resources and territoriality also play a role in the presence of horns in female bovids, but not
to the extent that defence against predators had (Stankowich & Caro, 2009).

Predation interactions play an important role in the evolution of horns in female bovids
(Abrams, 1990). It was found that Cheetahs will choose their prey based upon the energy
requirements necessary to catch the prey as well as the probability of injury (Clements, et al.,
2016). It is therefore beneficial for the individual to possess horns if they are an intermediate
size and cannot escape or take refuge as they will be better adapted to defend themselves
against predators (Clements, et al., 2016).

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In conclusion, the hypothesis that best explains the presence of horns in some female bovids
and not others, is the anti-predator defence hypothesis. This hypothesis is supported by
results that show the different drivers of horn evolution. For future studies it would be
necessary to investigate the relationship between fighting/intrasexual competition and the
reproductive success of female bovids as well as studying the reasons for different horn
shapes in female bovids and their significance.

Acknowledgements
I would like to thank Matthys Potgieter for his invaluable knowledge on the topic and Ronel
Nel for the guidance of my writing.

References
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