reef degradation and the loss of critical ecosystem goods and services provided by coral reef fishes

Reef degradation and the loss of critical ecosystem goods andservices provided by coral reef fishesMorgan S Pratchett1, Andrew S Hoey1 and Shaun K Wilson2,3

Available online at www.sciencedirect.com

ScienceDirect

Coral loss and declines in the physical structure of reef habitats

have marked effects on the abundance of coral reef fishes

which will be become even more pronounced as coral cover

continues to decline. Further, changes in the abundance and

composition of fish assemblages may have consequences for

ecosystem function and fisheries productivity. Even if fisheries

species are generally resilient to habitat degradation,

harvesting of larger fishes may threaten ecosystem functions

that are critical to coral reef resilience. Ultimately, managers will

need to explicitly address conflicts between increasing

demand for fish and the need to maintain ecosystem services.

Addresses1 ARC Centre of Excellence for Coral Reef Studies, James Cook

University, Townsville, QLD 4811, Australia2 Marine Science Program, Department of Parks and Wildlife,

Kensington, WA 6151, Australia3 Oceans Institute, University of Western Australia, Crawley, WA 6009,

Australia

Corresponding author: Pratchett, Morgan S

([email protected])

Current Opinion in Environmental Sustainability 2014, 7:37–43

This review comes from a themed issue on Environmental change

issues

Edited by Georgios Tsounis and Bernhard Riegl

For a complete overview see the Issue and the Editorial

Received 29 July 2013; Accepted 28 November 2013

Available online 21st December 2013

1877-3435/$ – see front matter, # Published by Elsevier B.V.

http://dx.doi.org/10.1016/j.cosust.2013.11.022

IntroductionHabitat degradation and habitat loss are the principal

causes of declining biodiversity [1], thereby jeopardizing

critical ecosystem goods and services. Habitat degra-

dation is largely manifest as loss of key habitat-forming

organisms (e.g. trees, kelp, and corals), which may be

replaced by smaller or less complex organisms, leading to

reduced habitat heterogeneity and structural (or topo-

graphic) complexity. Declines in habitat complexity

reduce the number and diversity of organisms that can

coexist, while fundamental shifts in habitat structure

often leads to localised extinctions of highly specialised

species. On coral reefs, scleractinian corals are major

contributors to both biological and physical habitat struc-

ture, and there is already evidence that sustained coral

loss in many geographical regions [2�,3] is leading to

www.sciencedirect.com

loss of biodiversity and erosion of important ecosystem

services [4,5�].

Coral reefs are among the most threatened ecosystems,

owing to a long-history of anthropogenic degradation and

exploitation [6], as well as disproportionate susceptibility

to emerging effects of global climate change [4,7]. Aside

from sustained coral loss, shifts in the biotic composition

of benthic habitats and declines in topographic complex-

ity are contributing to degradation of reef habitats

[2�,8�,9]. These changes are often accompanied by

declines in abundance of many reef-associated organisms,

especially coral reef fishes [10,11,12��] and non-coral

invertebrates [13].

The structural complexity of reef habitats is largely a

function of the abundance or cover of scleractinian corals

[14�]. However, coral cover and structural complexity may

vary independently when either the structural integrity of

erect coral skeletons is retained following extensive coral

mortality caused by bleaching or coral predators [10],

coral assemblages become dominated by growth forms

with low structural complexity [2�], or underlying com-

plexity is provided by geomorphic features, rather than

contemporary coral growth [15]. It generally appears that

loss of live coral independent of changes in habitat

structure have disproportionate effects on highly special-

ized fishes that live, feed, or settle on live corals [10,11],

whereas declines in topographic complexity may have

much broader impacts across a wide range of coral reef

fishes, including many of the larger reef fishes that sustain

tropical fisheries [16,50�].

Sustained and ongoing habitat degradation in coral reef

ecosystems will undoubtedly lead to the declines in

abundance of some reef fishes, but questions remain

about which species will be most affected and how the

structure and function of fish assemblages may change

with shifts in the biological and physical structure of

reef habitats. Of greatest importance is whether

ongoing coal loss and habitat degradation will lead to

the extinction of some species [17], undermine fisheries

productivity [5�], and/or erode key ecosystem functions

[18�,19�]. Using extensive data on the declines in

abundance of different coral reef fishes following acute

episodes of coral loss [12��], this study explores the

relative importance of first, body size, second, func-

tional roles of fishes, and third, ecological specialisation,

in explaining differential effects of reef degradation on

coral reef fishes.


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38 Environmental change issues

Figure 1

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Current Opinion in Environmental Sustainability

Alternative positive relationships between fish abundance and diversity

against coral cover and structural complexity. The widely assumed linear

relationship (a) suggests that habitat degradation will have similar effects

(in terms of proportional loss in abundance and diversity of fishes) no

matter what the absolute level of coral cover or complexity. The

threshold relationship (b) meanwhile would mean that the additional

effects of habitat degradation are most pronounced when coral cover

and complexity are already low.

Linear or thresholds effectsThe first step in establishing responses of fishes to coral

loss and habitat degradation is to establish the nature of

these relationships (Figure 1). The abundance of fishes

tends to increase linearly with increases in topographic

complexity measured, at least over the limited range of

values reported for fine-scale rugosity [14�]. It is also

assumed that abundance of fishes will increase in direct

proportion to live coral cover (Figure 1a). However, some

studies have failed to detect a significant relationship

between live coral cover and abundance of fishes, even

for species (e.g. coral-feeding butterflyfishes) with strong

reliance on corals [20]. These fishes may be insensitive to

changes in coral cover over the current range sampled, but

may decline in abundance following even more pro-

nounced coral loss.

Holbrook et al. [21] revealed non-linear relationships with

live coral cover for both abundance and diversity of reef

fishes (Figure 1b). These threshold responses suggest

that reef fishes will be generally resilient to changes in

live coral cover, so long as coral cover remains above 10–20%. Very large changes in coral cover (>20%) can,

however, have measurable effects on abundance of fishes,

and are particularly noticeable if coral loss is combined

with declines in topographic complexity [10,22]. More-

over, changes in total coral cover may belie changes in

coral composition, which can have equally important

effects for reef fishes that have strong reliance on particu-

lar types of corals [23].

The extent to which live coral cover regulates the

abundance of different fish will ultimately depend upon

the basis for their reliance on corals (e.g. food versus

settlement), their specificity in coral use, and the relative

importance of other regulatory processes (e.g. limited

larval supply). Reliance on live corals by coral reefs fishes

is often greatest at settlement, where up to 65% of

species settle amongst live coral [24]. This includes

some fishes that have limited dependence on corals as

adults [25,26] and some important fisheries target

species [27–29,30�]. Preferred corals for newly settled

fishes, as well as for many obligate coral-feeding and

coral-dwelling adult fishes, are erect branching species,

and predominantly Acropora [29,30�]. Further declines

in the abundance of corals (especially Acropora) will,

therefore, greatly constrain abundance of many different

reef fishes, especially if coral cover continues to decline

below 10–20% [3].

Selective effects of coral loss on reef fishesPrimary functional groups

Declines in the abundance of fishes that perform key

ecological functions (e.g. herbivores) are important in that

they may limit the capacity of reefs to recover following

coral loss [18�,31,32��]. If these fishes are negatively

affected by degradation of reef habitats, then this will


initiate a positive feedback that drives sustained and

accelerating degradation of reef habitats [18�]. Mainten-

ance of key ecological functions are often assumed based

on the high diversity of fishes within most primary func-

tional groups [33]. However, ongoing research is continu-

ing to unveil subtle differences in the ecological functions

of species placed within broad functional groups [18�,31].

Moreover, functional redundancy and the resilience of

coral reef ecosystems are critically dependant upon

response diversity among species with similar ecological

functions. Response diversity is rarely tested, but likely to

be low given that ecological function is often predicated

on strong reliance on specific resources [11].

Responses of fishes to coral loss and habitat degradation

do vary with trophic roles or functional groups [12��]. The

median response for all primary functional groups is

negative, though herbivores, carnivores and planktivores


Effects of reef degradation on coral reef fishes Pratchett, Hoey and Wilson 39

Figure 2

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Corallivores Carnivores Herbivores Planktivores

Res

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Response diversity apparent within primary functional groups of fishes, based on inter-specific differences in the responses of fishes to acute

disturbances that caused >10% declines in the cover or abundance of scleractinian corals. Data were compiled from multiple sources, as described in

Pratchett et al. [12��].

have higher response diversity compared to corallivores

(Figure 2), and some species within each of these groups

appear to benefit from coral loss. Importantly, marked

increases in abundance of some herbivorous fishes,

particularly grazing species [34�,35�], have been recorded

following extensive coral loss at some locations and are

probably critical to the prevention of phase shifts towards

macroalgal dominance [36]. However, response diversity

is substantially reduced when considering highly resolved

functional groups [12��]. The continued loss of live coral

is clearly going to lead to dramatic declines in the abun-

dance and diversity of corallivorous fishes and the func-

tions they perform. It remains to be seen, however, if

increases in a limited suite of species can offset the

extensive losses in other functional groups, thereby main-

taining key ecological processes as well as sustaining reef

fisheries. What is clear is that ongoing degradation of reef

habitats will further reduce functional redundancy [37]

making these systems even more vulnerable to future

disturbances.

Body size

Variation in body size and corresponding changes in the

scale of home ranges or feeding territories are key factors

responsible for response diversity among species with

similar ecological functions [38]. For coral reef fishes,

body size varies greatly [39] and is inextricably linked

to patterns of habitat use and the scale at which the

structure is provided. Importantly, live corals provide

relatively fine-scale complexity on coral reefs, benefiting

mostly small bodied and/or juvenile fishes [30�]. How-

ever, larger fishes, which play a disproportionate role in


key ecosystem processes [31,40] and contribute most to

fisheries catches, are expected to be less reliant on this

fine-scale complexity and relatively insensitive to coral

loss [15].

A prior meta-analysis that explored the extent to which

climate-induced coral bleaching compounds upon direct

threats to reef fishes from fisheries exploitation [41]

suggested that the large fishes targeted by reef fisheries

are generally immune from effects of coral loss and

habitat degradation. Alternative analyses suggest that

ongoing degradation of coral reefs and other coastal

habitats will certainly contribute to declines in pro-

ductivity (up to 20% by 2050) of tropical coastal fisheries

[5�,42], though effects of habitat degradation will be

compounded by several other more direct effects of

climate change on fishery species [43�]. Re-analysis of

the specific responses of 403 different coral reef species

[12��], comparing changes in abundance during acute

disturbances to species-specific records of maximum

length, revealed that mean responses were similar across

the full range of body sizes (Figure 3). Most of the fishes

that exhibited substantial declines in abundance were

small (20–50 cm in length). However, there were also

many small species that increased in abundance following

coral loss (with or without additional structural loss). For

larger fishes (>50 cm length), most species also declined

in abundance following coral loss, though some large

herbivorous and carnivorous species did exhibit increases

in abundance. The decline of larger bodied fish relates

partially to a higher dependence on live coral and associ-

ated structure during the early life history, when fish are



Figure 3

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(a) Coral loss only

(b) Coral loss and structural collapse

r2 = 0.001

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Relative change in the abundance of fishes ranging in size (reported maximum length) from 5 cm to 180 cm. Data based on changes in the abundance

of individual species during acute disturbance that caused (a) depletion of scleractinian corals without any changing in habitat structure, and (b)

declines in live coral cover that were associated with structural collapse of habitats, by virtue of the type of disturbance (e.g. cyclones) that caused

coral loss. Data was compiled from multiple sources, as described in Pratchett et al. [12��]. To see relationships specific to each primary function group

see Supplementary information.

smaller [27,28]. These data show that maximum achiev-

able body size alone does not explain variation in

responses of fishes to degradation of coral reef habitats,

and that large-bodied reef fishes, including targets of

many multi-species nearshore fisheries, will be vulnerable

to ongoing coral loss.


Ecological specialisation

The principal factor that has been shown to explain inter-

specific variation in responses of fishes to acute disturb-

ances is their ecological specialisation [11,44]. Not unex-

pectedly, highly specialised species with specific reliance

on a very small suite of different coral species, either for



food or habitat, experience greater declines in abundance

following extensive coral loss compared to generalist

counterparts that use a broader range of different corals

[44,45]. Severe or widespread depletion of specific coral

species may, therefore, lead to the extinction of highly

specialised reef fishes [17,45]. It is apparent, however,

that highly specialised species also have a number of

complementary traits that could offset their risk of extinc-

tion [17]. For example, highly specialised species often

dominate assemblages, reflective of the competitive

advantage conferred to specialists [46]. Even among

the most vulnerable fishes (e.g. Chaetodon trifascialisand other highly specialised coral feeding butterflyfishes)

there is some optimism for long-term persistence, given

high levels of gene flow apparent among widely dispersed

populations [17] such that it would require comprehen-

sive and simultaneous depletion of preferred prey (Acro-pora spp.) across virtually the entire Indo-Pacific to cause

global extinction of this species.

ConclusionsStrong associations between reef fishes and coral rich

habitats have been reported for many decades, though

the importance of these links has become particularly

apparent during recent and severe episodes of coral loss

[10,11,15,24]. Ongoing loss of live coral and associated

changes in the biological and physical structure of coral

reef habitats will unequivocally lead to declines in abun-

dance across a wide range of different coral reef fishes,

potentially undermining critical ecosystem processes,

and/or jeopardising sustainability of coral reef fisheries

[5�]. However, just as there are likely to be shifts in the

composition of coral assemblages, reflective of differen-

tial resilience among species [47�], there will be both

winners and losers among reef fishes [47�], leading to

changes in species composition. Highly specialised

species with a direct dependence on corals for food or

habitat are likely to be increasingly replaced with general-

ist species [48�]. The maintenance of ecosystem pro-

cesses, however, will depend on the functional

attributes/roles of the specific species that persist [49�].With the exception of corallivores, there appears to be

marked inter-specific variation in responses of species

within primary functional groups, providing some evi-

dence that high diversity and functional redundancy will

ensure continuity of ecosystem functions, at least during

moderate and short-lived declines of live coral cover. It

remains to be seen whether these increases in a limited

suite of species can offset reductions in the majority of

taxa in maintaining key ecological processes. Irrespective

of the compensatory potential of these species, reduced

diversity and hence functional redundancy within each

group will increase their vulnerability to future disturb-

ances and may lead to future ecological surprises.

The effects of coral loss and habitat degradation on the

overall productivity of reef-based fisheries are equivocal.


It is clear that there will be shifts in abundance and

composition of fishes with changes in habitat structure,

but in some instances these novel assemblages may

actually have similar or even higher biomass (albeit lower

diversity) of fishes that are harvested [8�]. Many of the

predominant species targeted by contemporary reef fish-

eries will decline in abundance following extensive coral

depletion [5�], either due to their inherent reliance on

coral habitats [27,28] or on prey that predominate in coral-

rich habitats [42]. Declines in abundance and diversity of

fishes will also become even more pronounced following

extensive coral loss and major shifts in the structure of

reef habitats. However, corresponding increases in the

abundance of some fishes may offset these declines,

leading to changes in catch composition, but limited

net loss of productivity. Even for fisheries species that

are generally resilient to habitat degradation, ongoing

harvesting of fishes may jeopardise ecosystem functions

leading to feedbacks that will further accelerate reef

degradation [18�]. Ultimately, managers will need to

explicitly address conflicts between increasing demand

for fish and the need to maintain ecosystem services,

which will become increasingly difficult within degraded

systems.

Appendix A. Supplementary dataSupplementary material related to this article can be

found, in the online version, at http://dx.doi.org/10.1016/

j.cosust.2013.11.022.

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Isolated reefs are often assumed to be highly vulnerable to large dis-turbance events owing to strong reliance on local replenishment, but alsobenefit isolation from most chronic anthropogenic disturbances (e.g.fisheries exploitation). This study documents the resilience of a small,isolated, offshore reef, in the aftermath of significant climate-inducedbleaching. Settlement and survivorship of corals were facilitated by localabundance of herbivorous fishes throughout.

36. Hughes TP, Rodrigues MJ, Bellwood DR, Ceccerelli D, Hoegh-Guldberg O, McCook L, Moltchaniwskyj N, Pratchett MS,Steneck RS, Willis BL: Phase shifts, herbivory and theresilience of coral reefs to climate change. Curr Biol 2007,17:360-365.

37. Hoey AS, Brandl SJ, Bellwood DR: Diet and cross-shelfdistribution of rabbitfishes (f. Siganidae) on the northern GreatBarrier Reef: implications for ecosystem function. Coral Reefs2013, 32:973-984.

38. Allen CR, Gunderson L, Johnson AR: The use of discontinuitiesand functional groups to assess relative resilience in complexsystems. Ecosystems 2005, 8:958-966.

39. Munday PL, Jones GP: The ecological implications of smallbody size among coral-reef fishes. Oceanogr Mar Biol Ann Rev1998, 36:373-411.

40. Bellwood DR, Hoey AS, Hughes TP: Human activity selectivelyimpacts the ecosystem roles of parrotfishes on coral reefs.Proc R Soc B 2012, 279:1621-1629.

41. Graham NAJ, Chabanet P, Evans RD, Jennings S, Letourneur T,MacNeil MA, McClanahan TR, Ohman MC, Polunin NVC,Wilson SK: Extinction vulnerability of coral reef fishes. Ecol Lett2011, 14:341-348.

42. Wilson SK, Fisher R, Pratchett MS, Graham NAJ, Dulvy NK,Turner RA, Cakacaka A, Polunin NVC: Habitat degradation andfishing effects on the size structure of coral reef fishcommunities. Ecol Appl 2010, 20:442-451.


43.�

Cheung WWL, Watson R, Pauly D: Signature of ocean warmingin global fisheries catch. Nature 2013, 497:365-369.

Using information on the temperature preferences of fish, combined withtemporal changes in fisheries catch data from around the world, thisstudy showed that global climate change and increasing ocean tempera-ture are affecting the composition, if not overall productivity, of temperatefisheries.

44. Pratchett MS, Coker DJ, Jones GP, Munday PL: Specialisation inhabitat use by coral reef damselfishes and their susceptibilityto habitat loss. Ecol Evol 2012, 2:2168-2180.

45. Munday PL: Habitat loss, resource specialization, and extinctionon coral reefs. Global Change Biol 2004, 10:1642-1647.

46. Blowes S, Pratchett MS, Connolly SR: Heterospecificaggression, dominance and specialization in a guild ofcorallivorous reef fishes. Am Nat 2013, 128:2-13.

47.�

Hughes TP, Baird AH, Dinsdale EA, Moltschaniwskyj NA,Pratchett MS, Tanner JE, Willis BL: Assembly rules of reef coralsare flexible along a climatic gradient. Curr Biol 2012, 22:736-741.

This study explores spatial variation in the abundance of different coraltaxa along a latitudinal, and therefore, temperature gradient, to infer likelychanges in the composition of coral assemblages as a consequence ofongoing ocean warming. Given marked differences in the distributionof corals it is concluded that climate change will lead to reassortment ofcoral reef taxa rather than wholesale loss of reef ecosystems.

48.�

Bellwood DR, Baird AH, Depczynski M, Gonzalez-Cabello A,Hoey AS, Lefevre C, Tanner JK: Coral recovery may not heraldthe return of fishes on damaged coral reefs. Oecologia 2012,170:567-573.

The diversity of small bodied crypto-benthic coral reef fish does notchange after coral bleaching; however, the composition of the fishcommunity becomes characterized by habitat generalists and there isa decline in coral specialists. These changes to fish community persist,even after coral recovery, implying that rapid changes in fish communitiesdue to coral loss are preserved for up to 13 years, even when specieshave short life-spans.

49.�

Cheal AJ, MacNeil MA, Cripps E, Emslie MJ, Jonker M,Schaffelke B, Sweatman H: Coral–macroalgal phase shifts orreef resilience: links with diversity and functional roles ofherbivorous fishes on the Great Barrier Reef. Coral Reefs 2010,29:1005-1015.

Comparing the trajectories of reefs following extensive coral mortality,this study highlighted the importance of the composition of herbivorousfish assemblages, with a shift to macroalgal-dominance being associatedwith low herbivore diversity, and low abundances of browsing andcropping species.

50.�

Wilson SK, Babcock RC, Fisher R, Holmes TH, Moore JAY,Thomson DP: Relative and combined effects of habitat andfishing on reef fish communities across a limited fishinggradient at Ningaloo. Mar Environ Res 2012, 81:1-11.

One of the few studies that explicitly assesses the interaction betweenhabitat condition on coral reefs and the effects of fishing, demonstratingthat the positive correlation between abundance of fish and structuralcomplexity is stronger on fished reefs than those where fishing is pro-hibited and that structural complexity is a better predictor of fish diversitythan live coral cover.

































































reef degradation and the loss of critical ecosystem goods and services provided by coral reef fishes

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