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Artificial Reefs, the Attraction-production Issue, and Density Dependence in

Marine Ornamental Fishes

Article  in  Aquarium Sciences and Conservation · March 2001

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Artificial reefs, the attraction-production issue, and
density dependence in marine ornamental fishes

JACQUELINE WILSON1 , CRAIG W. OSENBERG2 ,

COLETTE M. ST. MARY3 , CRAIG A. WATSON4 and
WILLIAM J. LINDBERG5
1,2,3
Department of Zoology, University of Florida, P.O. Box 118525, Gainesville,
FL 32611-8525, USA; 4 Department of Fisheries and Aquatic Sciences, University of Florida,
the Tropical Aquaculture Laboratory, Ruskin, FL 33570, USA; 5 Department of Fisheries and
Aquatic Sciences, University of Florida, Gainesville, FL 32653, USA

Key words: artificial reefs, density dependence, attraction, production, harvesting, marine
ornamentals

ABSTRACT

Artificial reefs may provide a useful tool to enhance production of marine ornamentals and to
divert detrimental harvesting activities from sensitive natural habitat. The efficacy of this strategy
depends, in part, on the extent to which artificial reefs contribute to new production (vs. attract
fishes from natural habitat) and therefore benefit harvested populations on a local and regional
basis. Here the attraction and production hypotheses and their application to marine ornamentals are
presented. Specifically, it is discussed how the strength and timing of density dependence can affect
the response of fish population dynamics to artificial reefs. In addition to this discussion, examples
of density dependence in marine ornamentals and related reef fishes are provided. Based on this
information, a simple conceptual model is presented to clarify the role of density dependence, and
this is followed by a discussion on the use of artificial reefs in the management and production of
marine ornamentals. Finally, unresolved scientific issues that remain to be addressed are provided.

INTRODUCTION

The vast majority of marine ornamental fishes sold in the aquarium trade are
collected from their natural habitats. Concerns have been raised about the environ-
mental impacts of these wild collections because harvesting may reduce the density
of marine ornamentals (via harvesting) and degrade habitat quality (via destructive
collecting methods: Jones and Steven, 1997; Jones and Hoegh-Guldberg, 1999).
One solution that would greatly reduce the environmental effects of the industry
is to increase the use of aquaculture and thus reduce the need for field collections.
Although this may be feasible for some species of fishes (e.g., anemone fish in
the genus Amphiprion, Bloch and Schneider, 1801), the majority (∼98%) of all
species sold in the aquarium trade cannot yet be produced in aquaculture (Martin
Moe of Green Turtle Publications, personal communication). As a result, harvest-
ing of ornamentals from the wild will be required to sustain the marine ornamental
industry, at least for the foreseeable future. Therefore, alternative harvesting and
management approaches that can facilitate environmentally sound and sustainable
exploitation of these marine resources are needed.

Aquarium Sciences and Conservation 3: 95–105, 2001.

© 2001 Kluwer Academic Publishers. Printed in the Netherlands.
96 J. WILSON ET AL.

One such scenario involves the use of artificial reefs. Artificial reefs are often
viewed as a fruitful way to not only increase fish production, but to also redirect
harmful human activities away from sensitive, natural reefs (Ambrose, 1994; Beets
and Hixon, 1994; Sosa-Cordero et al., 1998; Wilhelmesson et al., 1998). Although
the use of artificial reefs is typically focused on commercial and recreational food
fishes, they may also prove to be a useful, and as yet unexplored, tool in the man-
agement and augmentation of the marine ornamental industry. One vision (spear-
headed by Craig A. Watson and the Tropical Aquaculture Laboratory, Ruskin,
Florida) is to use specially designed artificial reefs to help facilitate the collec-
tion and production of marine ornamentals. For example, companies could lease
nearshore areas from a state and then deploy artificial reefs. They could then collect
fishes from these artificial reefs rather than from nearby natural reefs, which could
be protected from harvesting. This is similar to the way in which ‘live-rock’ is now
produced in Florida (Frakes and Watts, 1995). However, before such a tool can
be confidently embraced and successfully implemented, the ways in which artifi-
cial reefs can influence natural reef systems need to be evaluated more explicitly
(Osenberg et al., 2001).
Artificial reefs have not been discussed previously in the context of enhancing
marine ornamentals. They have, however, been discussed and used extensively for
other food and recreational fisheries (Beets and Hixon, 1994; Bortone et al., 1994;
Chua and Chou, 1994), but even in these contexts, there remains considerable
debate and uncertainty about their effectiveness. Before the use of artificial reefs
can be embraced, they must be more closely evaluated. This evaluation must include
a discussion that focuses on their effects on the population dynamics of natural
fish stocks, a topic that received little attention at the Marine Ornamentals’99
Conference in Kona, Hawaii, and which gave rise to this collection of papers.
The purpose of this paper is to discuss the conditions that influence the efficacy
of artificial reefs as management tools for harvesting marine ornamentals and to
introduce some fundamental scientific problems that must be addressed if artificial
reefs are to be used as management tools in the future. Specifically, a simple
conceptual model is proposed to address some of the controversy surrounding the
use of artificial reefs as management tools. The paper concludes with a discussion
of how an understanding of population dynamics and basic ecological processes,
such as density dependence, can help to determine the net effect of artificial reefs
on natural systems and the effect of harvesting on reef fish populations.

ATTRACTION VERSUS PRODUCTION

One of the central issues surrounding the effects of artificial reefs is the ‘Attraction
versus Production’ debate (Bohnsack, 1989; Lindberg, 1997). This debate is most
easily envisioned, and typically represented, as two opposing viewpoints. The
Attraction Hypothesis predicts that artificial reefs simply redistribute fishes without
augmenting production (Figure 1a). Although an artificial reef may have high
 ARTIFICI AL REEFS AND MARINE ORNAMENTAL FISHES 97

a) Attraction Hypothesis

Production Total

Artificial Reef

Natural Reef

Size of artificial reef

b) Production Hypothesis

Total
Production

Natural Reef

Artificial Reef

Size of artificial reef

Figure 1. Graphical representation of (a) the Attraction Hypothesis (in which total fish production
is constant), and (b) the Production Hypothesis (in which total fish production increases linearly
with the addition of new habitat).

densities of fishes, it does so because those fishes have been attracted away from
natural reefs. These fishes subsequently survive and grow at comparable rates on
artificial reefs as they would have on natural reefs. If artificial reefs only attract
fishes from other habitats, then as the size of the artificial reef increases, so will its
associated ‘production’ because more fishes will be pulled from the surrounding
natural reef habitat (Figure 1a). Conversely, the production associated with the
natural reef habitat will decrease with artificial reef size as more fishes leave the
natural habitat for the artificial reef. Therefore, although the spatial distribution of
fish production changes, total production associated with the natural and artificial
reef complex remains constant.
Fishes can be ‘attracted’ to artificial reefs at two different life stages – as larvae
(via settlement redirection) or as older age classes (via migration). Settlement is
the phenomenon when larval fishes leave the pelagic environment and settle to the
reef environment (see Keough and Downes, 1982), and it delineates the pelagic
98 J. WILSON ET AL.

from the benthic life stage. Because marine ornamentals are relatively small and
have high site fidelity, migration is likely to play a less important role than for
larger reef-dwelling fishes, which can move considerable distances after settlement
(but see Frederick, 1997). Therefore, settlement redirection will be concentrated
upon, and it will be assumed that migration of settled individuals is relatively
minor.
In contrast to the Attraction Hypothesis, the Production Hypothesis in this con-
text predicts that artificial reefs increase fish production by providing new habitat
to an otherwise saturated demersal environment. Thus, by providing new habitat
in the form of artificial reefs, larvae that would have otherwise never recruited
into older age-classes can instead settle and survive on the artificial reefs. Because
the artificial reef only produces new fish biomass rather than redistributes it, the
production associated with the artificial reefs will increase with increasing artifi-
cial reef size, but the production associated with the natural habitat will remain
constant. Therefore, there will be a net increase in the production of the overall
reef system (Figure 1b).
But what does this mean in terms of harvesting marine ornamentals? If artificial
reefs function according to the Production Hypothesis, artificial reefs could be
used to enhance the ornamental fishery as well as redirect potentially harmful col-
lections away from natural reefs. As a result, artificial reefs could be embraced as a
management tool without any significant detrimental effects to the dynamics of the
harvested fishes. However, if artificial reefs function according to the Attraction
Hypothesis and only redistribute fish biomass, then no new production is necessar-
ily associated with artificial reefs, and they represent a dangerous management tool.
In this case, deploying artificial reefs and attracting fishes away from the natural
reef habitat and then targeting these reefs with heavy harvesting for the ornamen-
tal industry would likely cause a decline in the overall fishery stock (Samples and
Sproul, 1985; Bohnsack, 1989). Thus, it is critical to understand how artificial
reefs affect the reef system in order to evaluate their use in management. Unfortu-
nately, there is little data with which to evaluate these hypotheses (Lindberg, 1997;
Seaman, 1997), in part because appropriate study designs have not been embraced
(Osenberg et al., 2001). Furthermore, artificial reefs probably both produce and
attract fish biomass – it is not useful to think of this as an ‘either/or’ issue.

EFFECT OF DENSITY DEPENDENCE ON THE RESPONSE OF

NATURAL REEFS TO ARTIFICIAL REEFS

Instead of taking a dichotomous approach, as often exemplified by the Attraction–

Production debate, an effective conceptual framework is needed. This framework
will allow for the quantification of the net production of a system, the partitioning
of that production to natural and artificial reef habitats and then the evaluation of
how much harvesting the aggregate system can likely sustain. The overall effect
of artificial reefs on fish population dynamics will depend, in part, on the degree
 ARTIFICI AL REEFS AND MARINE ORNAMENTAL FISHES 99

Relative production
on natural reef complete compensation

moderate d. d.

weak d. d.
no d.d.
Size of artificial reef
Figure 2. Graphical representation of how density dependence (d.d.) influences the effect of
artificial reefs on the production associated with a natural reef when there is attraction. Not shown
are cases when there is overcompensation (i.e., density dependence leading to increased production
on the natural reef: the line would lie above those shown) and depensation (i.e., facilitation: the
resulting line would lie below those shown).

of attraction as well as the strength of density dependence (Osenberg et al., 2001).

This is highlighted by considering a scenario in which attraction occurs, but the
strength of density dependence varies. The response of the natural reef to set-
tlement redirection changes as a function of the strength of density dependence
(Figure 2).
If fishes are attracted to the artificial reef and thus removed from the natural
reef, and there is no density dependence, there will be a decline in the production
associated with the natural reef (e.g., Figure 1a). If vital rates do not change with
density, then this results in the pure attraction scenario – production on the natural
reef would decline as a result of losing fishes to the artificial reef. If, however, certain
vital rates (e.g., settlement or per capita survival) increase in response to the initial
decline in density, then the production on the natural reef would be greater than
expected under the Attraction scenario (Figure 2). The greater the decline in density
(and the stronger density dependence is), the greater the compensatory response
would be. If the response in the vital rate completely compensates for the decline
in density (e.g., if settlement is reduced by 50% in response to the artificial reef, but
the survival of the remaining fishes is doubled because of the reduction in density),
then the density of older fishes would be unaffected by the artificial reef (Figure 2).
If density dependence is extremely strong, there could be ‘overcompensation’ and
the production of the natural reef could actually increase in response to the artificial
reef. If, conversely, survival or settlement decreases with decreasing density (i.e.,
exhibit depensation: Liermann and Hilborn, 1997), then the production of the
natural reef may be even less than predicted by the pure attraction scenario. So,
instead of one possible production curve for the natural reef (as depicted in the
Attraction scenario), there may be many, depending on the strength of density
dependence (Figure 2). Thus, even when there is Attraction, density dependence
could be sufficiently strong to produce a pattern that closely resembles the pure
100 J. WILSON ET AL.

Production Hypothesis (Figure 1b). By changing only the intensity of density

dependence, the Attraction and Production endpoints can both be predicted, as
well as all points in between (as well as some that are even more extreme) – net
production could increase, decrease or stay the same following the deployment
of an artificial reef. Thus, the efficacy of artificial reefs as a management tool
will depend critically on the strength of density dependence. How then is density
dependence produced?

DENSITY DEPENDENCE IN MARINE REEF FISHES

Most marine ornamental fishes have a bipartite life cycle (i.e., a dispersive, pelagic
larval stage followed by a relatively site-attached, benthic phase) (Sale, 1980;
Thresher, 1984). Density dependence can occur at any point in this complex life
history, but of interest in this context are the effects of density that arise at settle-
ment and later during the benthic stage – effects in the plankton are less relevant to
this discussion. Density dependence is used to refer to a negative (or positive) rela-
tionship between vital rates (settlement rates, or per capita post-settlement survival,
growth, or reproduction) and density. There are many mechanisms that produce
density dependence, including behavioral responses to chemical cues (e.g., in set-
ting settlement levels: Sweatman, 1988; Pawlik et al., 1991), social interactions
(e.g., via territoriality or mate-finding: Norman and Jones, 1984; Foster, 1985),
competition for food, shelter space or other critical resources (Hixon and Beets,
1993), and numerical and functional responses of predators (Murdoch and Bence,
1987).
To evaluate the importance of these processes to fish dynamics, it is critical to
distinguish between local (e.g., at the scale of a reef or < 1 km) and regional (e.g.,
> 100’s of km) dynamics. The local dynamics of a fish population are influenced
by factors that influence settlement rate and post-settlement survival. Because
eggs and larvae are dispersed via ocean currents (i.e., these local populations are
‘open’), the reproduction of adults on a local reef does not have a direct effect on the
local dynamics. This local reproduction does, however, contribute to the regional
production of larvae and is therefore critical in the sustainability of the overall
fishery. Growth of individual fish can have indirect effects on these demographic
rates by, for example, changing survival (via effects on size-dependent processes)
or by altering reproductive output (e.g., because fecundity often increases with
body size).
Although there is considerable debate about the strength of density dependence
in marine fishes (e.g., Caley et al., 1996; Steele, 1997; Hixon, 1998), there are
abundant data showing that it does operate and that it can occur at various points
in a fish’s life history. For example, the intensity of settlement (i.e., the number
of larval fishes that leave the pelagic environment and enter the reef environ-
ment) can be influenced by the number of older fish already present on the reef
(Figures 3a,b). In Coryphopterus glaucofraenum Gill, a small, short-lived goby
 ARTIFICI AL REEFS AND MARINE ORNAMENTAL FISHES 101

a) Coryphopterus glaucofraenum b) Dascyllus trimaculatus

4 100
Recruit density (no./m2)

Settler density (no./m2)

80
3

60
2
40

1 20

0
0
0 2 4 6 8 10 12
0 200 400 600 800 1000 1200
Adult density (no./m2)
Resident density (no./m2)

c) Coryphopterus glaucofraenum d) Pomacentrus ambionensis

1 35
length (mm) over 16 months
Change in mean standard
Proportion surviving

34
33
32
31
30
29
0.1 28
0 2 4 6 8 10 12 Low Med High
Initial density (no./m2) Juvenile density (no./patch reef)

Figure 3. Examples of density dependence in marine reef fishes. (a) Effects of resident adult
conspecifics on recruitment of the bridled goby, Coryphopterus glaucofraenum, in the Bahamas
(modified from Forrester, 1995). (b) Effects of resident conspecifics on the settlement of the
three-spot humbug, Dascyllus trimaculatus, in French Polynesia (modified from Schmitt and
Holbrook, 1996). (c) Effects of adult density on survival of adult Coryphopterus glaucofraenum
in the Bahamas over 2.5 months (modified from Forrester, 1995). (d) Effects of the density of
juvenile damselfish, Pomacentrus ambionensis (Bleeker, 1868), at One Tree Reef in Australia on
their growth over 16 months (low = 4 juveniles, medium = 8 juveniles, high = 12 juveniles,
see Jones, 1987 for more details). In (a) the number of recruits was assessed after 2.5 months
of cumulative settlement, and in (b) the number of settlers was assessed within 24 h of actual
settlement of larvae to the benthic habitat. In both (a) and (b) some post-settlement mortality
most likely occurred between initial settlement and subsequent assessment of recruits or settlers.
A Ricker Function (settler density = 0.5767∗[resident density]∗e(−0.0035)∗[resident density] ) was fitted to
the data in (b).

common in the Caribbean, settlement decreases as the density of resident adults

increases (Forrester, 1995). However, studies of other marine fishes have shown no
relationship between settlement and resident density (Steele et al., 1998), increas-
ing settlement with increasing adult density (i.e., positive density dependence
or ‘facilitation’; Sweatman, 1988), or hump-shaped relationships (Schmitt and
Holbrook, 1996: Figure 3b). Hence, not only the strength of density dependence,
but also the form of density dependence can vary. Later, the effect of density
dependence on harvesting strategies will be discussed.
102 J. WILSON ET AL.

Similarly, the density of resident fishes can also influence post-settlement sur-
vival. Forrester (1995) has shown that the subsequent survival of adult C. glau-
cofraenum decreases as the density of adults increases (Figure 3c). Thus, density
dependence can operate simultaneously at different points in the life history of a
single species. Density can also reduce growth due to competition for resources
(Figure 3d), and decreased growth can reduce reproductive rates because fecundity
is usually correlated with fish size. Although growth typically declines (or is not
affected) by increasing density, reproductive rates could initially increase with den-
sity via social or Allee affects (e.g., yielding depensation: Liermann and Hilborn,
1997) before declining due to the effects of competition. There are, however, little
data available on density dependent growth (but see Jones, 1987; Forrester, 1995;
Steele, 1998) and even less on density dependent reproduction. There are at least
two reasons for this. First, reproductive rates are difficult to measure (but growth
is not). Second, and more importantly perhaps, many researchers ignore density
dependent reproduction (and growth) in reef fishes given the open nature of these
systems – local reproductive output has no direct effect on the subsequent sup-
ply of larvae back to the local reef. This local larval production does, however,
affect ‘down current’ populations, and in order to understand the dynamics of
marine ornamentals over large spatial scales and manage the ornamental fishery
as a whole, better information regarding local reproduction and its link to larger
scale dynamics is needed.

ROLE OF DENSITY DEPENDENCE ON HARVESTING

Besides determining the net effect of artificial reefs on local production (Figure 2),
density dependence can also have important implications in terms of harvesting
(with or without respect to the use of artificial reefs): e.g., St. Mary et al. (2000).
To illustrate, consider the example of settlement using data for the three-spot
humbug damselfish, Dascyllus trimaculatus (Rüppell 1829), in French Polyne-
sia (Figure 3b: see Schmitt and Holbrook, 1996 for more details). In this sys-
tem, there are two opposing effects of density, which creates the hump-shaped
relationship: facilitation at low densities (i.e., residents provide chemical settle-
ment cues for new settlers), and inhibition at high density (i.e., mediated by com-
petition for shelter from predators) (Schmitt and Holbrook, 1996). As a result,
the highest settlement occurs at an intermediate density of conspecifics. This is
quite different from the pattern observed by Forrester (1995) for Coryphopterus
glaucofraenum (Figure 3a). All else being equal, the harvesting strategy for these
two species should also be quite different, owing to the different forms of density
dependence.

IMPLICATIONS FOR HARVESTING FISHES

For C. glaucofraenum, the greatest yield to a local collector targeting young fish
would be obtained by removing all residents, keeping the harvested species at low
 ARTIFICI AL REEFS AND MARINE ORNAMENTAL FISHES 103

local density, and thus maximizing the settlement rate. For D. trimaculatus, the
best strategy is different: the collector should keep residents at an intermediate
density in order to maximize settlement rates (this is a comparable situation to the
classic maximum sustained yield problem for closed populations). Hence, a very
high harvesting rate can actually reduce yield by greatly reducing resident density
and thus subsequent settlement rates.
These scenarios, however, focus on harvesting at a local scale, and ignore the har-
vesting that arises over much larger scales (involving many collectors in different
locales). For example, if C. glaucofraenum is depleted to low densities everywhere,
then the collecting rate at all locales will crash, owing to the lack of production of
larvae on a regional scale. Hence, the collection practices of a single collector may
be at odds with that of the entire industry (and the government agencies charged
with overseeing marine resources). This is one way in which artificial reefs may
help (but not necessarily). If artificial reefs produce fish, without any associated
impact on natural reefs, and if the deployment of artificial reefs is accompanied by
placement of natural reefs into a no-take reserve, then the local harvesting from
the artificial reef may be accomplished without any obvious negative impact to
the regional dynamics. If density dependence is sufficiently weak (or attraction
sufficiently strong) that the natural reef is negatively affected by the artificial reefs,
then this favorable scenario will not hold true.

FUTURE SCIENTIFIC NEEDS

The above discussion provides a very brief introduction to a very complex issue.
One hope is that it helps highlight the possible role of artificial reefs in the man-
agement and harvesting of marine ornamentals. More importantly, it is hoped that
this will help stimulate scientific research that can be used to better evaluate this
possible management tool. At present, there is insufficient information to lead to a
firm conclusion about the efficacy of artificial reefs. Indeed, there are many unre-
solved issues that prevent us from either embracing or rejecting the use of artificial
reefs as a management tool.
For example, the intensity of density dependence, how variable it is (among
species and environments), where it operates in the life history, and how it affects
the dynamics of reef fishes needs to be known. Although there has been consid-
erable experimental and observational work on these systems, there still is little
consensus about its occurrence and even less understanding of the patterns of
density dependence among different types of fishes, life stages or environmental
conditions. How artificial reefs might modify these processes on natural reefs must
be determined. In order to do this, the strength of density dependence operating
within the natural reef system must be quantified through experimental manipula-
tions and then how artificial reefs will modify these processes must be determined.
In so doing, the effect of artificial reefs on larval behavior during settlement and
on the depletion of the larval pool (i.e., via settlement redirection) must also be
104 J. WILSON ET AL.

understood. Such an understanding will require the integration of larval transport

mechanisms, larval behavior, and dispersal rates and distances. By learning more
about the basic ecology of this system (i.e., through experimental manipulation,
insightful monitoring, and adaptive management), managers and collectors alike
can begin to understand what drives particular processes of reef fish systems and
develop appropriate management scenarios and harvesting regimes.

ACKNOWLEDGEMENTS

We thank the Florida Sea Grant Program for their generous support for travel to the
Marine Ornamentals’99 Conference in Kona, Hawaii, and the College of Liberal
Arts and Sciences at the University of Florida for research support. We are also
grateful to Jennifer Piascik for her help with preparing graphics.

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Address for correspondence: J. Wilson, Department of Zoology, University of Florida, P.O. Box
118525, Gainesville, FL 32611-8525, USA
Phone: (352)392-9201, Fax: (352)392-3704; E-mail: [email protected]

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