The Factors Influencing Territorial Behaviour in

Herbivorous Fish


Emma Murrell-Orgill

S2846207



3803ENV Marine Biology




Date Submitted: 5
th
September 2014

Date Due: 5
th
September 2014



Word Count: 1866
Table of Contents
1.0 Introduction

2.0 Territorial behaviour
2.1 Food-related Territoriality
2.2 Site Attached Territoriality
2.3 Resting Site related Territoriality
2.4 Courtship related Territoriality

3.0 Non-Territorial behaviour
3.1 Schooling
3.2 Co-Existence
3.3 Spatial Distribution

4.0 Conclusion

5.0 References








1.0 Introduction
Within the Animal Kingdom, competition between species is a very common occurrence both
on land and underwater. Territoriality describes the competition for an area, as each
individual creates its own territory and defends it against any intruders, having specific
access to either food, mates and/or shelter. This behaviour has been observed amongst
herbivorous fish, however not all species compete. Species that have been reported to
compete include damselfish (Pomacentridae), surgeonfish (Acanthuridae), wrasses (Labridae)
and butterflyfish (Chaetodontidae) (Robertson and Gaines, 1986). A species ability to
economically defend an area (Brown, 1964) can be justified when the benefits of defending
the resource outweigh the energy costs of this behaviour, and are usually related to a limited
resource (Rooij et al., 1995). However research still needs to be conducted in order to
determine why some species still show territoriality when the resource they are defending is
not limited.
Herbivorous fish are found to display both territorial behaviour and non-territorial behaviour,
depending on a variety of factors. Some are able to successfully co-exist and share resources,
whereas others show a higher level of aggression to conspecific males and other species.
Some have overlapping territories like the damselfishes and surgeonfishes (Robertson and
Polunin, 1981), with the suggestion that this coexistence is able to occur through a symbiotic
and mutually beneficial relationship. This co-existence however rarely occurs within the
coral-reef systems, with a territorial species usually being present.
Research has also revealed that many of the reef fish are highly selective in respect to the
algae they consume (figure 1.), which may be associated with their unique morphology and
digestive capability when rupturing algal cell walls (Ogden and Lobel, 1978). Horn (1968)
found that territoriality was related to rich, stable food supplies where they can be
economically defended. This has led to the assumption that the more selective a fish is the
more territorial they are. Those that are not territorial are most likely to be less selective and
have a wide range of algal species they will consume.
Energy expenditure may also be a determining factor in respect to whether a species will
show aggression, as a higher amount of energy is needed to attack and chase away a
competitor. A higher amount of energy is required to successfully guard and chase away
competitors (Rooij et al., 1995), compared to those that are non-territorial.

Figure 1. List of different habitats and the associated fishes. (Source: Ogden and Lobel,
1978)
2.0 Territorial Behaviour
2.1 Food Related Territoriality
Brawley and Adey (1977) investigated the territorial behaviour of Threespot Damselfish, and
found that the males did exhibit this behaviour and excluded the smaller males along with the
Dusky Damselfish from Acropora cervicornis Lamarck patches, being their preferred food
source. Breeding also occurs within this male territory and coincides with the year-round
lunar cycle, with the females also laying their clutches here. It was found that the female
Threespots were also territorial, instead towards their preferred food source Montastrea
annularis. However contrary to this, research performed by Norman and Jones (1984) found
that there was no correlation found between the territory size and the abundance of algal food.
Instead it was perceived that intraspecific interactions refined the territory size, with the
addition of algal species to areas having no effect on territoriality, and only through the
removal of neighbours was there an observed change to the territory size.
2.2 Site Attached Territoriality
Research performed by Choat and Bellwood (1985) found that the surgeon fish Acanthurus
lineatus portrayed particular aggression towards other herbivorous fish which was site-
attached. They stated that
A.lineatus was distributed heterogeneously with high abundance at a single sample-
localityAbundances of herbivorous species at other localities did not correlate with
abundance patterns of A.lineatus.
However within the same report it was also found that they excluded species at some sites
and not others. When they researched further into this matter they found that the size of the
species was greater than standard length and therefore could affect the territoriality of the
A.lineatus.
2.3 Resting Site Related Territoriality
Robertson and Sheldon (1979) investigated the relationship between wrasse Thalassoma
bifasciatum, and two small territorial damselfish Eupomacentrus dorsopunicans and E.
planifrons in regards to their sleeping shelter habitat. Damselfish are only territorial towards
wrasse in respect to their food and eggs, not sleeping sites. They described the wrasse to
regularly change their holes, with little intra- or interspecific aggressive interaction. When
its hole is removed, a wrasse is late in retiring but finds a hole near its old one with little
aggressive interaction, and does not have a higher mortality rate.
Robertson and Sheldon found that the behaviour exhibited by the wrasse was performed due
to the risk of predation increasing when returning to their hole is delayed, however sleeping
sites were found to be in high abundance so therefore further research needs to be conducted
in order to determine the hidden factors leading to their sleeping site related territoriality.
2.4 Courtship related territoriality
Male Cichlid fish Pseudosimochromis curvifrons were found to display different behaviours
depending on the species. Research conducted by Kuwamura (1992) revealed that the males
attacked conspecific males and other species, but only chased the conspecific males out of
their territory. The other species were only attacked when they intruded the spawning sites
where the males court or wait for the females at one site, compared to other sites where they
were attacked while patrolling and foraging.

3.0 Non-territorial
3.1 Schooling
Instead of defending a territory, Robertson et al. (1976) observed that the non-territorial
Striped Parrotfish Scarus Croicensis were attacked less by territory owners whilst as a school,
compared to those non-schooling. By doing this, schooling allows individuals who are
unable to successfully obtain a feeding territory to bypass their competitors in a simple,
energy efficient way. Higher energy input is required to chase away a competitor so therefore
the S. croicensis has overcame this by utilising defence in numbers.
3.2 Co-existence
Herbivorous fish species have been found to co-exist and essentially benefit through a
mutualistic relationship. van Rooij et al., (1995) observed this behaviour in the stoplight
parrotfish, Sparisoma viride, suggesting that
factors that may favour territory sharing between herbivores are fine-scale resource
partitioning and shared defence, both of which would reduce the costs of territorial life.
This behaviour has also been observed by Robertson and Sheldon (1979) between wrasse and
territorial damselfish, however this only occurs when it is time to retire to their holes.
Damselfish have the ability to defend the holes and prevent the wrasse from entering the
holes, increasing their chance of predation and therefore minimising competition, but instead
they portray little to no aggression.
3.3 Spatial Distribution
When investigating the different zones, Bruggemann et al (1994) observed the territorial
single males of the stoplight parrotfish Sparisoma viride within the deeper reef zones,
compared to large numbers of non-territorial fish amongst the shallow reef (figure 2).

Figure 2. Diagram showing the relationship between zones and behaviour of
herbivorous reef fish. (Source: Bruggermann et al., 1994).
They concluded that foraging of individuals in restricted to certain reef areas as a result of
social structure of S.viride.
Bruggemann et al., (1994) investigated whether these parrotfish really need to delve deeper
levels in order to access the superior food sources, as it is unknown as to whether or not
territoriality can be deemed a practical approach or even necessary. They found that S.viride
grows continuously throughout its life and engages in reproductive activities throughout the
year. These activities suggest that protein and energy may constrain individual performance
confirming that they have specific energy needs, found to grow below the lower limits of the
their territorial areas (Bruggemann et al, 1994). S.viride are territorial to an area that does not
include their preferred food source, requiring further research to determine the factors
preventing them from moving their territory to the area of interest.
4.0 Conclusion
Literature has shown that herbivorous fish are an extremely diverse and remarkable group
with the ability to be either territorial or not, and still successfully survive and reproduce.
This presents the idea that it is neither a benefit nor detriment to be territorial, and that there
is other ways to overcome this without high-energy expenditure. There is still uncertainty as
to what makes a herbivore territorial, whether it be their taxonomic status (Bruggemann et al.,
1994) or highly-selective food requirements (Horn, 1984). Herbivores can be territorial when
other species or conspecific individuals are competition for food resources (Brawley and
Adey, (1977) or during the breeding season (Kuwamura 1992), or they have site-attached
aggression (Robertson and Sheldon 1979; Choat and Bellwood 1985). Some herbivorous fish
chose to swim in schools (Robertson et al. 1976) instead of being territorial, which appears to
be the most energy-efficient and successful approach to gaining access to their preferred
feeding sites. Species tend to be territorial or not, and there is only a couple of species that
have successfully found a way to mitigate aggressive species. However there is still no clear,
defining evidence to support why a species is territorial whilst others are not.

5.0 References
Brawley, S. H., & Adey, W. H. (1977). Territorial behavior of threespot damselfish
(Eupomacentrus planifrons) increases reef algal biomass and productivity. Environmental
Biology of Fishes, 2(1), 45-51.
Bruggemann, J. H., van Oppen, H., & Breeman, M. (1994). Foraging by the stoplight
parrotfish Sparisoma viride. I. Food selection in different, socially.Marine Ecology Progress
Series, 106, 41-55.
Choat, J. H., & Bellwood, D. R. (1985). Interactions amongst herbivorous fishes on a coral
reef: influence of spatial variation. Marine biology, 89(3), 221-234.
Kuwamura, T. (1992). Overlapping territories ofPseudosimochromis curvifrons males and
other herbivorous cichlid fishes in Lake Tanganyika. Ecological Research, 7(1), 43-53.
Lobel, P. S. (1980). Herbivory by damselfishes and their role in coral reef community
ecology. Bulletin of Marine Science, 30(Supplement 1), 273-289.
Norman, M. D., & Jones, G. P. (1984). Determinants of territory size in the pomacentrid reef
fish, Parma victoriae. Oecologia, 61(1), 60-69.
Ogden, J. C., & Lobel, P. S. (1978). The role of herbivorous fishes and urchins in coral reef
communities. Environmental biology of fishes, 3(1), 49-63.
Robertson, D. R., & Sheldon, J. M. (1979). Competitive interactions and the availability of
sleeping sites for a diurnal coral reef fish. Journal of Experimental Marine Biology and
Ecology, 40(3), 285-298.
Robertson, D. R., Sweatman, H. P. A., Fletcher, E. A., & Cleland, M. G. (1976). Schooling as
a mechanism for circumventing the territoriality of competitors. Ecology, 1208-1220.
Sale, P. F. (1976). The effect of territorial adult pomacentrid fishes on the recruitment and
survival of juveniles on patches of coral rubble. Journal of Experimental Marine Biology and
Ecology, 24(3), 297-306.
van Rooij, J. M., de Jong, E., Vaandrager, F., & Videler, J. J. (1996). Resource and habitat
sharing by the stoplight parrotfish, Sparisoma viride, a Caribbean reef
herbivore. Environmental Biology of Fishes, 47(1), 81-91.