Coral Reefs (2009) 28:683–690

DOI 10.1007/s00338-009-0501-0

REVIEW

Herbivory versus corallivory: are parrotfish good or bad

for Caribbean coral reefs?
Peter J. Mumby

Received: 19 February 2009 / Accepted: 14 April 2009 / Published online: 3 May 2009
Ó Springer-Verlag 2009

Abstract With coral cover in decline on many Caribbean parrotfishes in facilitating coral recruitment, growth, and
reefs, any process of coral mortality is of potential concern. fecundity. In contrast, no net deleterious effects of coral-
While sparisomid parrotfishes are major grazers of Carib- livory have been reported for reef corals. Corallivory is
bean reefs and help control algal blooms, the fact that they unlikely to constrain overall coral cover but contraints
also undertake corallivory has prompted some to question upon dwindling populations of the Montastraea annularis
the rationale for their conservation. Here the weight of species complex are feasible and the role of parrotfishes as
evidence for beneficial effects of parrotfishes, in terms of a vector of coral disease requires evaluation. However, any
reducing algal cover and facilitating demographic pro- assertion that conservation practices should guard against
cesses in corals, and the deleterious effects of parrotfishes protecting corallivorous parrotfishes appears to be unwar-
in terms of causing coral mortality and chronic stress, are ranted at this stage.
reviewed. While elevated parrotfish density will likely
increase the predation rate upon juvenile corals, the net Keywords Conservation
Mortality
Coral
Herbivore
effect appears to be positive in enhancing coral recruitment
through removal of macroalgal competitors. Parrotfish
corallivory can cause modest partial colony mortality in the Introduction
most intensively grazed species of Montastraea but the
generation and healing of bite scars appear to be in near Parrotfish are some of the most abundant fish on Caribbean
equilibrium, even when coral cover is low. Whole colony reefs and they usually dominate the biomass of herbivorous
mortality in adult corals can lead to complete exclusion of fishes ([80%), which also includes acanthurids, kyphosids
some delicate, lagoonal species of Porites from forereef and pomacentrids. The impacts of parrotfish feeding have
environments but is only reported for one reef species been studied since the 1960s (Randall 1965) and while it
(Porites astreoides), for one habitat (backreef), and with differs among species (Bruggemann et al. 1996; Burkepile
uncertain incidence (though likely \\10%). No deleteri- and Hay 2008), the vast majority of feeding activity is
ous effects of predation on coral growth or fecundity have directed towards algal substrates, including encrusting
been reported, though recovery of zooxanthellae after corallines, algal turfs and macroalgae (Barlow 1975; Gygi
bleaching events may be retarded. The balance of evidence 1975; Bruggemann et al. 1994a). Not surprisingly, exper-
to date finds strong support for the herbivory role of imental and observational studies have found that parrot-
fish grazing can exert strong top–down impacts on the
cover of macroalgae on reefs (Hay 1981; Carpenter 1986;
Communicated by Biology Editor Dr. Philip Munday Lewis 1986; Steneck 1997; Williams and Polunin 2000;
Williams et al. 2001; Mumby et al. 2006; Burkepile and
P. J. Mumby (&) Hay 2008). Given concern over the deleterious conse-
Marine Spatial Ecology Lab, School of BioSciences, Hatherly
quences of phase shifts towards increased macroalgal cover
Laboratory, University of Exeter, Prince of Wales Road, Exeter
EX4 4PS, UK (Done et al. 1996), many authors have argued that parrot-
e-mail: [email protected] fish conservation should, in principle, benefit coral reefs

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and the ecosystem services they provide (Steneck 1988;

Hughes 1994; Bellwood et al. 2004; Newman et al. 2006;
Hoegh-Guldberg et al. 2007; Knowlton and Jackson 2008;
Mumby and Steneck 2008; Nystrom et al. 2008). The point
made by these studies is that the grazing behaviour of
parrotfish should benefit corals by helping to limit their
algal competitors.
While parrotfishes primarily feed from algal substrates,
corallivory has also been widely reported (Fig. 1), partic-
ularly from species in the genus Sparisoma such as S. vi-
ride and S. aurofrenatum (Barlow 1975; Frydl 1979;
Bruggemann et al. 1994b; Bruckner and Bruckner 1998;
Bruckner et al. 2000; Sanchez et al. 2004; Rotjan and
Lewis 2006). Two forms of parrotfish corallivory have
been described; ‘spot biting’ in which individual bite-sized
lesions are distributed over the coral skeleton and ‘focused
biting’ in which a larger, continuous patch of coral tissue is
excavated by repeated, overlapping predation (Fig. 2;
Bruckner et al. 2000).
With coral cover having declined in much of the
Caribbean (Gardner et al. 2003), any process of corallivory
has the potential to exacerbate other sources of coral
mortality such as coral disease or bleaching (Kramer 2003).
Indeed, the dual trophic role of parrotfishes in grazing
algae and corals has led some to question the rationale for
conserving corallivorous parrotfishes (Rotjan and Lewis
2008). This review aims to assess the net impact of

Fig. 2 Bite patterns of parrotfish corallivory showing a spot biting on

Porites astreoides and b focused-biting on Colpophyllia natans.
Photo credit: Andrew Bruckner

parrotfish feeding on the population dynamics of corals.

Given that corals underpin ecosystem services, processes
of corallivory are considered to be negative impacts
whereas the consumption of algae is considered a positive
effect. In short, the review attempts to weigh the evidence
for negative and positive impacts of parrotfish on Carib-
bean reefs, and in so doing, highlights implications for reef
management and future research. The impacts of parrotfish
grazing are reviewed separately for each major demo-
graphic process of corals.

Coral recruitment

The degree to which parrotfishes prey directly or inadver-

Fig. 1 Predation of Montastraea annularis by a terminal phase tently on coral recruits is unknown. In a study of macro-
stoplight parrotfish, Sparisoma viride. Photo credit: Andrew Bruckner algal impacts on recruits of the coral Agaricia (diameter

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1 cm), Box and Mumby (2007) found that 15% of uncaged reserves in the Caribbean, Mumby et al. (2007a) found a
corals suddenly disappeared. They attributed this mortality strong positive correlation (r [ 0.8, P \ 0.001) between
to parrotfish predation rather than physical dislodgement the grazing activity of the parrotfish community and the
by wave action because none of the caged treatments density of juvenile corals (diameter up to 2 cm). Coral-
exhibited such sudden mortality, and it was unlikely that livorous species of parrotfish dominated the community yet
caging effects would have been sufficient to account for a coral recruitment increased with grazing, largely in
refuge from wave disturbance in all replicates given the response to the decrease in macroalgae. The study dis-
high flow in some microhabitats that were not potentially counted potentially confounding effects of reef rugosity,
sheltered by adult colonies. Moreover, Bak and Engel also damselfish abundance and coral cover on coral recruitment,
observed that grazing fishes could damage juvenile corals and used high-resolution models of potential larval supply
(Bak and Engel 1979). In contrast, Birkeland (1977) found to discount the likelihood that the pattern of grazing
no evidence of predation upon coral recruits, when exam- coincided with that of larval supply. In short, available
ining coral settlement substrates after they had been grazed evidence suggests that parrotfish grazing has a net positive
intensively by acanthurids and small parrotfishes in Pan- impact on coral recruitment.
ama. While it should be borne in mind that Box and
Mumby (2007) did not prove that parrotfish predation was Partial colony mortality
the cause of recruit mortality, the two studies may be partly
reconciled by considering the parrotfish species involved. Some of the most detailed studies of partial colony mor-
The study by Box and Mumby (2007) was carried out on a tality in Caribbean corals took place before the late 1990s
forereef in Honduras and S. viride, possibly the most (Hughes and Jackson 1985; Bythell et al. 1993; Meesters
important parrotfish corallivore (Miller and Hay 1998), et al. 1996; Meesters et al. 1997), after which coral diseases
dominated the parrotfish community (Box 2008). The have increasingly become a major source of partial mor-
predominant parrotfish grazers in Birkeland’s (1977) study tality (Weil et al. 2006). Parrotfish corallivory was found to
were small parrotfishes, primarily of the species Sca- be an important source of chronic partial colony mortality
rus iserti, which rarely feeds on coral (Mumby, personal in studies from the US Virgin Islands (Bythell et al. 1993)
observation). and Curaçao (Meesters et al. 1996; Meesters et al. 1997).
Parrotfish grazing has been hypothesized to facilitate Not surprisingly, the number of bite scars on corals has
coral recruitment through at least three mechanisms. been found to increase with colony size (Meesters et al.
Firstly, grazing of the reef surface helps promote encrust- 1997), though, whether this indicates a preferential selec-
ing coralline algae and limit macroalgae, thereby increas- tion of larger colonies by parrotfishes or merely reflects the
ing the settlement space available for coral planulae null expectation that bite prevalence will scale positively
(Steneck 1988). Secondly, grazing helps prevent the with colony surface area is not clear.
establishment of thick algal turfs which can trap sediment Not all coral species are equally susceptible to parrotfish
and induce post-settlement mortality in coral recruits corallivory. In those reef habitats where they are present,
(Birkeland 1977; Birrell et al. 2005). Similarly, by reducing the coral species Montastraea faveolata and M. annularis
the cover of macroalgae, grazing may reduce the frequency tend to have the greatest prevalence of bite scars, often
and duration of competitive interactions between coral exceeding 20% of colonies (Garzon-Ferreira and Reyes-
recruits and macroalgae. Contact between coral recruits (at Nivia 2001). Bythell et al. (1993) found that up to 10% of
least of the genus Agaricia) and common macroalgae such the tissue surface area of M. annularis could have been
as Lobophora variegata and Dictyota pulchella can reduce removed by parrotfish in St. Croix. In contrast, several
recruit growth rates dramatically (Box and Mumby 2007), coral species including Montastraea cavernosa, Diplo-
and possibly reduce their survivorship because the time ria strigosa and adult encrusting corals like Agari-
spent at vulnerable, small sizes is extended. cia agaricites show very low prevalences of corallivory,
Overall, if corallivorous species of parrotfish are com- often less than 1% (Bythell et al. 1993; Garzon-Ferreira
mon on a reef, it seems reasonable to expect a degree of and Reyes-Nivia 2001; Reyes-Nivia et al. 2003). The coral
parrotfish-based mortality on coral recruits. Logically, if Porites astreoides appears to experience intermediate lev-
the number of corallivorous parrotfishes increases on a els of predation (Bythell et al. 1993). In Belize, Rotjan and
reef, then the predation rate on coral recruits would be Lewis (2005) generally found low levels of predation on
expected to increase also. So, is there any evidence that this species, and while the prevalence of scarred colonies
increasing the number of parrotfish actually has a net varied among reef zones, the greatest prevalence reported
negative impact on coral recruitment? Apparently not. In a was only 4%, and often either 2% or absent. However, in
study of the impact of the Exumas Land and Sea Park the minority of corals that exhibited predation, the intensity
which is one of the oldest and most successful marine of grazing could be great with nearly half (48%) of the

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scarred colonies in backreef areas having lost at least half shallow areas). Rotjan and Lewis (2005) categorised 10%
of their tissue. of P. astreoides colonies in a Belizean backreef system as
Despite the relatively high predation pressure on exhibiting whole colony mortality due to parrotfish pre-
M. annularis, bite scars usually heal within 1-2 months dation. While parrotfish predation may indeed have been
(Bythell et al. 1993; Sanchez et al. 2004). In a particularly the cause of mortality, a causal link was not verified by
revealing study, Sanchez et al. (2004) transplanted 23 ramets observation, and it remains possible that alternative pro-
of M. annularis to a highly-grazed forereef at 10 m. At the cesses caused the mortality and that parrotfish grazing
beginning of the study, all ramets were free of grazing scars. happened to be intense on the subsequent dead substrate.
For the following 13 months, a monthly census was Uncertainty over the actual causation notwithstanding, it
undertaken of the extent of parrotfish-induced mortality appears that whole colony mortality in reef-associated
on colonies, and used to parameterise a transition matrix coral species is—at worst—confined to a small proportion
of four injury states: 0% of tissue consumed, \ 1% of colonies, within a single species (P. astreoides), and a
consumed, \ 5% consumed, [ 5% consumed. The study single reef habitat (backreef). Even on these corals, the
revealed that processes of injury (predation) and recovery are demographic significance of such mortality is unclear
remarkably dynamic. By the end of the study all ramets had because dead corals can remain in an intensively grazed
experienced injury at some stage but 50% were free of any state for months (Rotjan and Lewis 2005), if not years.
bite scars, and only a small fraction (\ 10%) of ramets had Thus, the actual mortality rate might be considerably lower
more than 5% of missing tissue. In general, the modal state of than the observed prevalence of dead, yet intensively
a ramet was to remain with \ 1% injury (probability 0.31 per grazed, corals.
month) and the transitions between no bite scars and \ 1%
injury were virtually at equilibrium (probability of 0%
to \ 1% was 0.17 and from \ 1% to 0% was 0.16). Coral growth, physiology, and reproduction
In summary, parrotfish corallivory varies significantly
among species with M. annularis and M. faveolata being Chronic forms of stress, including coral bleaching, have
among the most intensively-consumed species and Diplora been found to exert deleterious impacts on coral growth
strigosa and A. agaricites being among the least. However, (Mascarelli and Bunkley-Williams 1999). Whether par-
even the most intensively-grazed species appears to have a rotfish corallivory exerts a significant deleterious influence
high capacity for tissue regeneration and there is little on the physiology of the coral holobiont remains unclear.
evidence that processes of predation exceed recovery even There is clearly a metabolic cost to healing bite lesions but
on small ramets of coral. no evidence to date that such costs extend to significant
reductions in coral growth rate or fecundity. A study of
Whole colony mortality (of post-recruits) zooxanthellae densities in the coral Montastraea spp. found
that recovery rates shortly after a minor bleaching event
The effects of parrotfish predation in causing whole colony (and hurricane) were lower in colonies that had experi-
coral mortality appears to be restricted to a few, highly- enced some parrotfish predation ([6 scars) than in colonies
palatable species. Working in the Florida Keys, Miller and that had no grazing scars (Rotjan et al. 2006). Whether
Hay (1998) found that delicate colonies of Porites div- differences in the zooxanthellae communities between
aracata were completely consumed by sparisomids within grazed and non-grazed colonies persist for long after the
48 hours of being transplanted from seagrass beds to the bleaching event is unclear, as are the physiological impli-
forereef. Similarly, intense predation by S. viride has cations of such differences in symbionts for coral growth,
excluded the highly-palatable, small branching coral Por- but the study implies that bite lesions may be a contributory
ites porites furcata from parts of a Belizean backreef that form of stress in Montastraea.
are deep enough to allow access for larger herbivores The direct effects (if any) of parrotfish predation on the
(Littler et al. 1989). Another species of Porites, P. ast- fecundity of coral colonies has not yet been quantified.
reoides has a harder skeleton (Littler et al. 1989) and, while However, contact between common Caribbean macroalgae,
experiencing significant predation (Rotjan and Lewis including Dictyota and Lobophora, during the period of
2005), is not excluded from either forereef or backreef gametogenesis in the coral M. annularis have been shown
habitats (Littler et al. 1989; Miller and Hay 1998). to reduce egg size, the number of eggs per gonad and the
Few, if any, Caribbean cases of parrotfish-driven whole number of gonads per polyp (Foster et al. 2008). Thus, the
colony mortality have been described for corals that are removal of macroalgae by grazing is likely to alleviate
typically found in reef habitats for which large-bodied macroalgal impacts on coral fecundity and help maintain
fishes have access (i.e., all reef habitats except very larval connectivity.

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Discussion 2007), reduce the growth rate of adult P. astreoides

(Chasqui-Velasco et al. 2007) and P. porites (Miller and
The title of this article, ‘Are parrotfish good or bad for Hay 1998) and reduce the fecundity of M. annularis (Foster
Caribbean reefs?’, is intended to be provocative rather than et al. 2008). Comparable results have also been found in
pose an appropriate question for discussion; few aspects of the Pacific (Tanner 1995; Jompa and McCook 2002;
nature warrant such polarised viewpoints. The real aim of Hughes et al. 2007). Clearly, the positive impacts of
this review is to weigh the negative demographic impacts grazing are unequivocal, although the full demographic
of corallivory against the positive demographic impacts on implications for corals after sustained reductions in mac-
coral of reducing the cover of algal competitors. roalgal cover have not yet been measured in the field.
Parrotfish corallivory has received a modest degree of Balancing the positive effects of grazing and negative
research over the last two decades and the conclusions of impacts of corallivory on coral population dynamics is
most studies appear to be consistent. Evidence for negative challenging and warrants continued study. One study that
influences of corallivory principally find that corallivory is explored this question directly for Porites porites con-
able to exclude a few highly palatable species of Porites cluded that the negative impact of greater corallivory on
from reefs and that predation-based partial colony mortal- coral growth was balanced by the positive impact of
ity can be modest (10% of tissue) in the most intensively grazers removing algal competitors (Miller and Hay 1998).
grazed species of Montastraea, but its prevalence is gen- However, even here, while corallivory reduced coral
erally low (\4%) in the community for most species. growth rate, coral growth remained positive. To date, no
Claims that ‘parrotfish predation has a direct adverse effect study has demonstrated net negative impacts of corallivory
on coral survival’ (Rotjan and Lewis 2005) are not borne on coral population dynamics. Of course, the critical
out by data at a population scale. As a source of partial question is whether this scenario might change in future.
coral mortality, bite scars on the most intensively grazed From the perspective of total coral cover which supports
corals can heal within weeks, generating a potentially many ecosystem services (Done et al. 1996) and has the
equilibrial dynamic between injury and recovery (Sanchez desirable property of enhancing grazing intensity and
et al. 2004). Corallivory may constitute a source of acute reducing macroalgal cover (Williams et al. 2001), it is
mortality in coral recruits, but the available evidence difficult to conceive that corallivory will cause a net neg-
implies that any negative impacts are outweighed by ative impact. The reason for this is that many of the coral
positive effects in removing algal competitors (Mumby species that recruit and grow rapidly on Caribbean reefs,
et al. 2007a). With the exception of a few (\1%) inten- such as Agaricia spp, are relatively lightly affected by
sively grazed coral colonies, there is little evidence that parrotfish predation. Indeed, some of these ‘weedy’ spe-
parrotfish corallivory contributes to whole colony mortality cies, such as Agaricia, are particularly vulnerable to mac-
in adult corals. One possible exception is P. astreoides in roalgal overgrowth (Nugues and Bak 2006) and are
backreef habitats, but even here where whole colony therefore poised to benefit significantly from a reduction in
mortality has been attributed to corallivory (Rotjan and macroalgal cover in response to elevated parrotfish
Lewis 2005), the demographic significance of such mor- biomass.
tality is likely to be limited. For example, if we assume that While corallivory might not be expected to affect total
heavily grazed corals can remain in that state for 2 years coral cover significantly, the story could be different for the
(after which they become colonised by fleshy algae), then major framework-builders M. faveolata, M. franksi, and
the observed prevalence of 10% of P. astreoides colonies M. annularis. Natural processes of recruitment appear to be
found in a wholly grazed state (Rotjan and Lewis 2005) highly sporadic in these species (Edmunds and Elahi 2007),
could, in principle, reflect an underlying parrotfish-driven and asexual reproduction appears to be a significant mode
coral mortality rate of only 5% or less (assuming that the of reproduction in at least M. annularis (Foster et al. 2007).
system is in steady state). Such low levels of mortality are It is unknown whether rates of corallivory on juveniles of
predicted to have limited demographic consequences for Montastraea are exceptionally high, but some species in
this rapidly recruiting species (Mumby 2006). this genus are susceptible to a recruitment bottleneck
From a coral perspective, the positive effects of grazing (Edmunds 2002; Mumby 2006). With the continued rise in
in reducing the cover of thick algal turfs and macroalgae disease incidence on these species (Weil et al. 2006), the
have been reviewed elsewhere (Burkepile and Hay 2006; future persistence of Montastraea is questionable in many
Mumby and Steneck 2008). In summary, thick algal turfs locations. Of particular concern is whether the incidence of
and macroalgae can kill juvenile corals (Birkeland 1977), corallivory will increase as coral cover declines and whe-
cause partial colony mortality of some adult corals (Lewis ther this might lead to demographic consequences for these
1986; Lirman 2001; Nugues and Bak 2006), cause a ces- long-lived corals that appear to have limited potential for
sation of growth in juvenile A. agaricites (Box and Mumby recovery. The evidence to date is limited but does not

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imply a problem. For example, the study of corallivory on fully understood (Bruggemann et al. 1994a) and while
M. annularis by Sanchez et al. (2004) implicitly created a parrotfishes prefer to feed from lower density coral sub-
scenario of low coral cover in a high grazing environment strates (Bruggemann et al. 1996), the reasons for favouring
by placing corals within a zone of dead Acropora. The particular coral species or even individuals are unclear.
authors found no evidence for net deleterious impacts on Indeed, the underlying reasons for corallivory remain
corals. speculative and include the marking of territorial bound-
Further insight into the effects of corallivory at low aries. The response of parrotfishes to fluctuating avail-
coral cover can be obtained from ecological models of ability of coral is also uncertain; for example, how does the
coral and algal dynamics and empirical studies of reef rate of coral predation scale with parrotfish density and the
recovery. The question can be stated explicitly as, ‘does availability of prey? At low coral cover, will predation tend
corallivory prevent coral recovery once total coral cover to be intensive on individual coral colonies—tending to
becomes low?’ An initial ecological model of Caribbean cause whole colony mortality—or extensive, tending to
coral population dynamics used a parameterisation for cause chronic partial colony mortality on a larger number
partial colony mortality from the study by Bythell et al. of colonies? Similarly, will parrotfishes feed on a greater
(1993), in which parrotfish grazing was a major component diversity of coral prey as the abundance of preferred spe-
of the observed mortality (Mumby 2006). Despite such cies declines? Importantly, future research might discover
corallivory, coral recovery was found to occur from low whether corallivory can exert an additive or synergistic
cover providing that grazing levels were high enough to affect on coral growth with other stressors, such as algal
remove an adequate amount of algae. The model parame- competition or reduced aragonite saturation state. The role
terisation for corallivory was extended in a follow-up study of parrotfishes as vectors of coral disease also needs to be
by adding an explicit parrotfish predation rate on coral investigated given that other corallivores have been asso-
recruits (from Box and Mumby 2007) and observed rela- ciated with the transmission of some coral diseases (Wil-
tionships from Curaçao (Meesters et al. 1997) for the liams and Miller 2005). With many uncertainties
incidence of parrotfish lesions with colony size and extent notwithstanding, any assertion that conservation practices
of partial mortality on such colonies (Mumby et al. 2007b). should guard against protecting corallivorous parrotfishes
While this model did not increase the rate of corallivory (Rotjan and Lewis 2008) appears to be unwarranted at this
above that observed in situ, coral recovery was found to be stage.
feasible despite what are likely to be relatively high levels
of corallivory, given the density of large herbivores in the Acknowledgements I thank the Marine Spatial Ecology Lab for
stimulating discussions on this topic and the helpful comments of
sites used to parameterise the model. Laurence McCook and two anonymous referees. Andy Bruckner
While models predict that corallivory should not retard kindly provided photographs to illustrate the article.
coral recovery if grazing levels are high, there are few (if
any) tests of this prediction. Few monitoring studies have
observed low coral cover during a period of high grazing. References
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