See

discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/228032221

Niche-assembly vs. dispersal-assembly rules

in coastal fish metacommunities: Implications
for...

Article in Journal of Applied Ecology May 2007

DOI: 10.1111/j.1365-2664.2007.01330.x

CITATIONS READS

32 118

1 author:

David Mouillot
Universit de Montpellier
334 PUBLICATIONS 10,959 CITATIONS

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Soil macroinvertebrate functional traits View project

Liteau PAMPA View project

All content following this page was uploaded by David Mouillot on 20 February 2014.

The user has requested enhancement of the downloaded file.

Journal of Applied
Niche-assembly vs. dispersal-assembly rules in coastal fish
Blackwell Publishing Ltd

Ecology 2007
44, 760767 metacommunities: implications for management of
biodiversity in brackish lagoons
DAVID MOUILLOT
UMR CNRS-UMII 5119 Ecosystmes Lagunaires, Universit Montpellier II, CC 093, 34095 Montpellier cedex 5,
France

Summary
1. Biodiversity is rapidly being lost in a world transformed increasingly by human
activities. We need to determine urgently the factors which control the coexistence of
species and thus allow local biodiversity to be maintained.
2. Coexistence between interacting species can be explained by species-sorting, mass-
effect, patch dynamic and neutral perspectives on metacommunities. According to the
first two paradigms, community assembly rules are based on species ecological niches
or functional roles, whereas the latter two emphasize the role of stochastic processes.
3. Here I consider whether effective strategies for managing fish biodiversity in coastal
lagoons depend on the predominant paradigm.
4. The neutral perspective was not considered relevant because lagoon fishes belong to
different trophic levels and have clear niche differences.
5. Where fish coexistence is ruled by species-sorting, priority must be given to the
preservation or restoration of habitat, whereas heterogeneity among lagoons and
population densities become critical when mass effects predominate.
6. According to the patch dynamics perspective, the key factors are individual
turn-over and the fitness equivalence among species.
7. Synthesis and applications. Coastal fish metacommunities are not ruled consistently
by a single theoretical paradigm. However, consideration of such extreme cases sets
clear boundaries for species assembly rules. Confronting the three main views is not
only a controversial topic, but also has consequences for ecosystem and biodiversity
management.
Key-words: fishing pressure, habitat preservation, heterogeneity, invasion, niche, species
coexistence
Journal of Applied Ecology (2007) 44, 760767
doi: 10.1111/j.1365-2664.2007.01330.x

the processes underpinning local losses remain unclear

Introduction
Acceleration of biodiversity loss is recognized widely
in both terrestrial (Thomas et al. 2004) and marine
ecosystems (Roberts & Hawkins 1999) and, in aquatic
ecosystems, the most important factors are certainly
climatic change and biotic exchanges (Roberts & Hawkins
1999; Nystrom, Folke & Moberg 2000). Although the
causes of biodiversity loss seem clear at a global scale,
2007 British
Ecological Society
Correspondence: David Mouillot, UMR CNRS-UMII 5119
No claim to original Ecosystmes Lagunaires, Universit Montpellier II, CC 093,
French government 34095 Montpellier cedex 5, France (fax: +33 467143 719; e-
works mail: mouillot@University-montp2.fr).
(Mouquet & Loreau 2003). Although the consequences
of changes in local biodiversity have led to some debate
(Kinzig, Pacala & Tilman 2002), this may be reaching a
consensus (Loreau et al. 2001; Hooper et al. 2005), and
it is clear that preserving biodiversity has both direct
and indirect benefits.
A key element in understanding biodiversity and
how it changes is to understand the factors that
determine how species coexist. The coexistence of
species is explained classically by either a niche-
assembly or a dispersal-controlled perspective on
communities. The unified neutral theory of biodiversity
and biogeography (Hubbell 2001) assumes the per capita
761 ecological equivalence of individuals (whatever the lagoon ecosystem. Secondly, the more fish species there
Biodiversity species they belong to) and this dispersal-related view are in the coastal lagoons, the more various trophic
management in contrasts with proposals that coexistence is maintained levels can be expected to exist, increasing the complexity
coastal lagoon fish by niche differentiation. In niche-assembled communities, of the food web. Thirdly, the more fish species there are
communities interacting species are at equilibrium with, according in the coastal lagoons, the more varied the biological
to the limiting similarity theory (MacArthur & Levins responses can be expected to be during a perturbation
1967), the best competitor occupying each ecological or a crisis event, increasing the resistance of the system.
niche. These two views of species coexistence are not Finally, the more fish species there are in the coastal
mutually exclusive and are actually under scrutiny with lagoons, the more ecological niches can be occupied,
regard to their ability to explain the relative abundance increasing the resistance to invasion (Stachowicz et al.
of species (McGill 2003; Volkov et al. 2003) and to predict 2002).
species invasion (Fargione, Brown & Tilman 2003; Hence, the processes by which the number of coex-
Herben et al. 2004). isting fish species can be maintained is of major concern
Much of the formal theory about community ecology for coastal lagoons. My aim is to consider whether
focuses on a single scale, but local communities are not effective strategies for managing fish biodiversity in
isolated assemblages and it is now generally accepted coastal lagoons depend on which metacommunity
that community ecology has to be investigated within a paradigm predominates and, if so, to propose appropriate
metacommunity framework (Hubbell 1997; Loreau, management priorities.
Mouquet & Gonzalez 2003; Leibold et al. 2004). A
metacommunity is defined as a set of local communities
Coastal lagoon fish and the metacommunity
that are linked by dispersal of multiple, potentially
framework
interacting species (Wilson 1992). Of the four paradigms
proposed in metacommunity theory (see review in Metacommunity models have either been based on a
Leibold et al. 2004), three assume niche differences among local community connected to a regional pool or on a
species and one proposes neutrality among individuals. set of spatially assembled local communities. In the
Here I propose to fill the gap between these theoretical former, the local community interacts with the regional
metacommunity paradigms and practical implications pool (the metacommunity) through migration (Bell 2001;
for management of biodiversity in brackish lagoons. Hubbell 2001) while, in the latter, the metacommunity
Coastal lagoons provide essential ecosystem services is partitioned into spatially explicit local assemblages
such as shoreline protection, water quality improvement that are connected by individual migrations (Chave,
and fisheries resources, as well as habitat and food for Muller-Landau & Levin 2002). A coastal lagoon
migratory and resident animals and recreational areas comprises a set of localities (patches of macrophyte,
for human populations (Levin et al. 2001), and represent macroalgal, sandy, muddy, shellfish or rocky habitat),
13% of the world coastline. These highly productive which hold local communities and are connected to
ecosystems (300 g C m2 year1) (Knoppers 1994) are each other to constitute a lagoon metacommunity (Fig. 1).
under severe stress: in addition to climatic change
inducing sea-level rise, human impacts have led to
problems such as anoxia, over-exploitation of resources,
destruction of habitats, eutrophication and pollutant
contamination from land use in the watershed (Joyeux
& Ward 1998; Crooks & Turner 1999).
Many of the goods and services provided by coastal
lagoon ecosystems rely on the species inhabiting these
areas, with fish influencing ecosystem processes through
trophic relationships with other compartments (e.g.
Mancinelli, Costantini & Rossi 2002). Within lagoon
fish communities we can distinguish freshwater (occa-
sional visitors via freshwater discharges), brackish-
water and marine species. Most of the marine species
are transients (or at least non-residents), using lagoons
as either feeding or breeding areas (Levin et al. 2001)
and therefore exporting biomass and, indirectly,
nutrients to adjacent marine ecosystems. Fish may also
2007 British accumulate and export pollutants (Mouillot et al.
Ecological Society
2000). The study of fish diversity is important for
No claim to original
French government
several reasons. First, the more fish species there are in
Fig. 1. The regional fish metacommunity in coastal
works, Journal of the coastal lagoons, the more body shapes, colours or ecosystems whereby the lagoon fish metacommunity (which,
Applied Ecology, 44, swimming abilities can be observed (after recreational in turn, contains local communities associated with particular
760767 fishing or not), increasing the aesthetic value of the localities) is connected to the regional pool of fish.
762 The fish metacommunity within a lagoon is connected
D. Mouillot to a larger-scale, regional metacommunity, embracing
all coastal fish populations potentially able to colonize
the lagoon. Metacommunity paradigms will be ex-
amined here at the regional level: spatial dynamics
occurring among habitat patches within the lagoon
metacommunity will not be discussed.
It is clear that fish species are not strictly equivalent,
either in coastal lagoons (e.g. Dumay et al. 2004) or in
coastal ecosystems in general (e.g. Bellwood et al. 2006
found a high functional versatility among coral reef fish
belonging to the same trophic group). Thus the neutral
theory is not relevant at the scale of biological or functional
heterogeneity seen in coastal fish and the neutral met-
acommunity perspective will not be considered further.
Three paradigms might shape coexistence of fish
species in coastal lagoons (Fig. 2). The species-sorting
perspective postulates that environmental heterogeneity
among and within localities is strong enough to promote
spatial niche segregation among species. Strong gradients
in abiotic factors or in habitat characteristics (three
niches in Fig. 2a) lead to local environmental conditions
acting as a niche filter on the regional pool of species
(Zobel 1997; Statzner, Doledec & Hugueny 2004).
2007 British
Ecological Society
No claim to original
French government Fig. 2. From metacommunity perspectives to management strategy for promoting lagoon fish richness. Solid arrows indicate
works, Journal of higher dispersal rates between lagoons and the sea than dashed arrows. The extent to which a species is the competitive dominant
Applied Ecology, 44, in a lagoon habitat is given by the matching of the species symbol (denoting its habitat niche type) with the habitat symbol. Smaller
760767 symbols for species indicate smaller-sized populations.
Local community composition is thus fairly resilient to
disturbance (Cottenie et al. 2003), and coexistence of
species in the metacommunity is ruled by niche parti-
tioning processes (e.g. Albrecht & Gotelli 2001). The
mass-effect perspective (Fig. 2b) postulates that
poor competitors can escape from local competitive
exclusion by mass immigration. As in the species-
sorting perspective, environmental differences among
localities are strong enough to provide spatially segre-
gated niches, but these localities are sufficiently con-
nected by dispersal to establish sourcesink relations
and thus to allow species (such as species C in Fig. 2b)
to colonize localities where they are poor competitors
from adjacent localities where they are better com-
petitors and are thus massively represented (Gonzalez
et al. 1998; Mouquet & Loreau 2003). The third, patch
dynamics perspective, postulates that localities are
identical and that species coexistence is maintained by
a trade-off between competitive ability and dispersal
(Levin & Paine 1974). All species have the same
probability of being present in a locality: even if species
A is the strongest competitor, species B and C can
also be present in the lagoon because of their better
dispersal ability (Fig. 2c).
763
From metacommunity paradigms to a biodiversity
Biodiversity
management strategy
management in
coastal lagoon fish Here I consider the priorities for managing fish
communities biodiversity in coastal brackish lagoons from the
metacommunity viewpoint. Although these three pure
theoretical rules are an oversimplification, their use has
the merit of producing clear hypotheses and is poten-
tially valuable for proposing management priorities.
According to both the two first perspectives (Fig. 2a,b),
spatial niche differentiation occurs among localities
within the lagoon (Fig. 1) but fish diversity within the
lagoon is maintained by different processes (the
presence of species C is promoted either by the presence
of a niche where it is one of the strongest competitors or
is dependent on a high regional density).
the species-sorting perspective
According to the species-sorting perspective, the number
of species is limited by the number of niches available
within the lagoon. Consequently, a large spectrum of
alpha (resource-related) or beta (habitat-related) niches
is needed to maintain or increase fish richness (Wilson
1999). The number of beta niches depends on both
diversity and heterogeneity of habitats (Ziv 1998;
Guidetti et al. 2002) and, in coastal lagoons, habitat
diversity is often related to macrophyte diversity and
cover (Sfriso, Birkemeyer & Ghetti 2001), with macrophyte
beds and meadows providing microhabitats and
protection from predators (Dolbeth et al. 2003).
Transplanted macrophyte beds can provide the
structural and functional attributes of natural beds in
previously degraded sites (Dolbeth et al. 2003) and
mobile macrofauna often colonize the new habitat
within a couple of weeks (Sogard 1989). Fish and shrimp
richness was therefore higher in transplanted eelgrass
beds than on unplanted substrates after less than 1 year
(Fonseca et al. 1990). Shellfish aquaculture can also
provide new habitats, with oyster reefs, for instance,
creating corridors between shelter and foraging grounds
(Peterson & Lipcius 2003; Peterson, Grabowski &
Powers 2003). Sea urchins also provide biogenic struc-
tures and thus new microhabitats that allow more fish
species to become established (Hartney & Grorud 2002).
In order to increase the number of alpha niches avail-
able in coastal lagoons we can increase the diversity of
resources and the diversity of prey. Resource partitioning
is a key factor in promoting fish coexistence because
competition is reduced when diet overlap is low (DePirro,
Marchetti & Chelazzi 1999; Sibbing & Nagelkerke 2001;
Carrasson & Cartes 2002). The amount and diversity of
food sources are again dependent on the heterogeneity
2007 British and diversity of habitats, with oyster reefs, for instance,
Ecological Society
providing structured habitat for finfish, crabs and other
No claim to original
French government
organisms (Breitburg et al. 2000) and other man-made
works, Journal of structure providing suitable habitats for amphipods in
Applied Ecology, 44, brackish lagoons (Aikins & Kikuchi 2001). Nevertheless,
760767 Parker, Duffy & Orth (2001) demonstrated experimentally
that most indices of animal diversity in marine en-
vironments were more strongly related to total vegetal
surface area than to vegetal diversity, indicating that
increasing vegetation cover is another strategy for
maintaining a large alpha niche spectrum in brackish
lagoons where fish communities are niche-assembled.
The diversity of both beta and alpha niches depends
on water quality. Eutrophication and its associated
oxygen imbalance (Justic 1991) and increasing organic
particulate matter ratio (DellAnno et al. 2002) has
become a major problem in numerous coastal areas
(Flindt et al. 1999). Environmental constraints such as
oxygen depletion (Ishitobi et al. 2000) are likely to
reduce the number of fish species because fewer species
are able to tolerate eutrophic conditions (e.g. Ludsin
et al. 2001). Increased primary production may trans-
late into additional production for pelagic fishes but
not for demersal ones feeding on benthic resources. Also,
periodic dystrophic crises observed in coastal lagoons
with high eutrophication levels (Bachelet et al. 2000)
lead to high mortality of both fish and invertebrates
and to a degradation of fish habitats (Karim, Sekine &
Ukita 2003; Powers et al. 2005; Bishop et al. 2006) and,
inevitably, to a decrease in species richness. Watershed
management must therefore aim to decrease nutrient
and pollutant inputs (Ludsin et al. 2001). Dispersion is
also critical for maintaining local diversity; because
species must be able to reach the niches they are able to
occupy, sea-lagoon exchanges must be carefully
maintained.
In summary, when richness of the fish community
within a lagoon depends on species-sorting, management
should mainly focus on habitat protection, habitat
diversification within the lagoons, quality improvement
of run-off waters and biotic exchanges (Fig. 2).
the mass-effect perspective
If mass-effects operate, a species can be present in a
lagoon even if it is not a good competitor in any of the
available niches, providing the species (C in Fig. 2b) is
abundant somewhere that can act as a source of
individuals for the sink lagoon. Management priorities
are therefore rather different: they need to preserve a
high regional density for all fish populations, to maximize
dispersal and thus maintain connectivity between
habitats, and to promote regional habitat diversity to
ensure that each species has a habitat where it is among
the best competitors.
Strong heterogeneity among coastal habitats will
prevent any biotic homogenization at the regional
scale (Olden et al. 2004) and thus preserve the source
sink relationships and the mass effects (Mouquet &
Loreau 2003). In highly eutrophic lagoons, planktiv-
orous and detritivorous fish (low trophic levels) are
likely to maintain dense populations, whereas carniv-
orous demersal fishes will benefit from a high second-
ary production of invertebrates in more oligotrophic
lagoons where they will reach higher densities. Different
764 lagoons may act as complementary sources of various
D. Mouillot populations, each one providing optimal conditions
for a given group of species (Fig. 2b).
the patch dynamics perspective
In the patch dynamics view, local habitats within the
lagoon are considered to be equivalent (Fig. 2c). Fish
species coexistence is permitted by trade-offs between
functional attributes and is ruled by the dispersal-
assembly hypothesis, i.e. communities are non-
equilibrium assemblages of species, their presence or
absence being dictated by random dispersal. A fish
species that is a poor competitor can be present in the
lagoon if it compensates with high dispersal abilities
(such as species C in Fig. 2). In this case we need to
preserve the dispersal of fish individuals, for instance
by managing channels to allow fish movements
throughout the year and reducing or removing passive
nets and other physical barriers (Crespi 2002). Fish
dispersal and migration are also strongly influenced
by physical oceanographical features such as shifts in
hydrological conditions in coastal lagoons and estuaries
(Garcia et al. 2004). Fish can detect a wide range of
external stimuli (chemicals, temperature and atmospheric
pressure) (Bullen & Carlson 2003) and rapid shifts in
hydrological conditions are likely to modify routes and
intensity of fish migration (Brehmer et al. 2006) and,
thus, richness under this perspective.
Variability of abiotic factors within the lagoon will
promote turn-over in species composition, suggesting
that lagoons must be protected from increasing marine
influence (and thus more stable temperature, salinity
and pH). A hydrological shift from hypohaline (late
1950s) to hyperhaline conditions in the Terminos
lagoon (Gulf of Mexico) restricted the richness, the
taxonomic diversity and density of estuarine species, as
well as the presence of freshwater species following
freshwater discharge from the watershed (Ramos
Miranda et al. 2005; Sosa-Lopez et al. 2007).
Another critical point is to preserve the fitness
equivalence among species within the regional pool.
Functional differentiation of the mosquitofish Gambusia
affinis (Baird & Girard) (Dumay et al. 2004), its higher
aggressiveness than native species (Mills, Rader & Belk
2004) and its greater dispersal abilities than some
congeners (Rehage & Sih 2004) all contribute to the
relative invasiveness of this species. Particular attention
must therefore be paid to invasive species which may
break the trade-off rule and thus prevent coexistence
and richness of fish species within coastal lagoons.
2007 British Conclusion
Ecological Society
The current management programmes for biodiversity
No claim to original
French government
are largely pattern-based and focus on hotspots (Turpie,
works, Journal of Beckley & Katua 2000; Roberts et al. 2002). They
Applied Ecology, 44, should become process-based and include functional
760767 mechanisms. Thus, preserving goods and services
provided by ecosystems requires not only the investi-
gation of biodiversity patterns but also the identification
of processes that generate and maintain local biodiversity:
different metacommunity paradigms lead to different
management strategies to maintain lagoon fish bio-
diversity. Priorities for increasing local richness in coastal
lagoon fish communities and the number of functions
provided by the species depend upon the communities
assembly rules.
Although it is unlikely that any coastal fish metacom-
munity is ruled solely by one of the three hypotheses,
one assembly rule may be predominant. Indeed,
Leibold & Norberg (2004) suggested that plankton
metacommunities may be ruled by species-sorting
paradigm. Regional fish metacommunities embracing
lagoon and coastal habitats may be consistent with the
mass-effect paradigm because the many different patches
enable niche segregation (Mouillot, Dumay & Tomasini
2007), connectivity among coastal sites is very high
(Sanvicente-Anorve, Flores-Coto & Chiappa-Carrara
2000) and because regional habitat diversity (e.g.
Mouillot et al. 2005) may promote sourcesink rela-
tionships between localities. In the open sea, where
differences among localities are much less pronounced,
fish metacommunities are more likely to follow the
patch dynamics paradigm.
The challenge now is to move from conjecture and
speculation to confront metacommunity paradigms
with empirical data. Assembly rules in lagoon fish can
be investigated using functional differences to test
whether coexisting species in local patches are more
similar than expected by chance (i.e. than random
samples from the regional pool). If so, local habitat
characteristics act as environmental filters, allowing
only a narrow spectrum of species to survive (Peres-
Neto 2004; Mouillot et al. 2007) and species-sorting
and mass-effect are better explanations than patch-
dynamics. If coexisting species are less similar than
expected by chance (i.e. compared to random samples
from the pool of species able to survive in these
conditions) we may conclude that interspecific
competition limits species similarity (Jensen 1997) even
in connected habitats, and that species-sorting is the
best candidate. Innovative approaches, using acoustic
methods (Gaudreau & Boisclair 2000), might allow
estimation of fish fluxes between coastal lagoons and
the sea (dispersal) and evaluation of fish density within
the lagoon (carrying capacity). In any case, before
proposing an integrated management strategy for
local biodiversity in limnetic ecosystems, it is necessary
to take into account the processes that rule local
assemblages in metacommunities.
Acknowledgements
I wish to thank J. C. Joyeux, T. Bouvier and J. A.
Tomasini for comments on the manuscript. J. B. Wilson
and R. de Wit corrected the English and provided
useful suggestions.
765
References
Biodiversity
management in
Aikins, S. & Kikuchi, E. (2001) Studies on habitat selection by
coastal lagoon fish amphipods using artificial substrates within an estuarine
communities environment. Hydrobiologia, 457, 7786.
Albrecht, M. & Gotelli, N.J. (2001) Spatial and temporal
niche partitioning in grassland ants. Oecologia, 126, 134
141.
Bachelet, G., de Montaudouin, X., Auby, I. & Labourg, P.J.
(2000) Seasonal changes in macrophyte and macrozoobenthos
assemblages in three coastal lagoons under varying degrees
of eutrophication. ICES Journal of Marine Science, 57,
1495 1506.
Bell, G. (2001) Neutral macroecology. Science, 293, 24132418.
Bellwood, D.R., Wainwright, P.C., Fulton, C.J. & Hoey, A.S.
(2006) Functional versatility supports coral reef biodiversity.
Proceedings of the Royal Society of London, Series B,
Biological Sciences, 273, 101107.
Bishop, M.J., Powers, S.P., Porter, H.J. & Peterson, C.H.
(2006) Benthic biological effects of seasonal hypoxia in a
eutrophic estuary predate rapid coastal development.
Estuarine, Coastal and Shelf Science, 70, 415 422.
Brehmer, P., Do Chi, T. & Mouillot, D. (2006) Amphidromous
fish school migration revealed by combining fixed sonar
monitoring (horizontal beaming) with fishing data. Journal
of Experimental Marine Biology and Ecology, 334, 139150.
Breitburg, D.L., Coen, L.D., Luckenbach, M.W., Mann, R.,
Posey, M. & Wesson, J.A. (2000) Oyster reef restoration:
convergence of harvest and conservation strategies. Journal
of Shellfish Research, 19, 371377.
Bullen, C.R. & Carlson, T.J. (2003) Non-physical fish barrier
systems: their development and potential applications to
marine ranching. Reviews in Fish Biology and Fisheries, 13,
201212.
Carrasson, M. & Cartes, J.E. (2002) Trophic relationships in a
Mediterranean deep-sea fish community: partition of food
resources, dietary overlap and connections within the
benthic boundary layer. Marine Ecology Progress Series,
241, 4155.
Chave, J., Muller-Landau, H.C. & Levin, S.A. (2002) Comparing
classical community models: theoretical consequences for
patterns of diversity. American Naturalist, 159, 123.
Cottenie, K., Michels, E., Nuytten, N. & De Meester, L.
(2003) Zooplankton metacommunity structure: regional
vs. local processes in highly interconnected ponds. Ecology,
84, 9911000.
Crespi, V. (2002) Recent evolution of the fishing exploitation
in the Thau lagoon, France. Fisheries Management and
Ecology, 9, 1929.
Crooks, S. & Turner, R.K. (1999) Integrated coastal management:
sustaining estuarine natural resources. Advances in Ecological
Research, 29, 241289.
DellAnno, A., Mei, M.L., Pusceddu, A. & Danovaro, R.
(2002) Assessing the trophic state and eutrophication of
coastal marine systems: a new approach based on the
biochemical composition of sediment organic matter.
Marine Pollution Bulletin, 44, 611 622.
DePirro, M., Marchetti, G.M. & Chelazzi, G. (1999) Foraging
interactions among three benthic fish in a Posidonia oceanica
reef lagoon along the Tyrrhenian Coast. Journal of Fish
Biology, 54, 1300 1309.
Dolbeth, M., Pardal, M.A., Lillebo, A.I., Azeiteiro, U. &
2007 British Marques, J.C. (2003) Short- and long-term effects of
eutrophication on the secondary production of an intertidal
Ecological Society
macrobenthic community. Marine Biology, 143, 1229
No claim to original
1238.
French government
Dumay, O., Tari, P.S., Tomasini, J.A. & Mouillot, D. (2004)
works, Journal of
Functional groups of lagoon fish species in Languedoc
Applied Ecology, 44, Roussillon, southern France. Journal of Fish Biology, 64,
760767 970983.
Fargione, J., Brown, C.S. & Tilman, D. (2003) Community
assembly and invasion: an experimental test of neutral
versus niche processes. Proceedings of the National Academy
of Sciences USA, 100, 89168920.
Flindt, M.R., Pardal, J.A., Lillebo, A.I., Martins, I. &
Marques, J.C. (1999) Nutrient cycling and plant dynamics
in estuaries: a brief review. Acta Oecologica, 20, 237248.
Fonseca, M.S., Kenworthy, W.J., Colby, D.R., Rittmaster,
K.A. & Thayer, G.W. (1990) Comparison of fauna among
natural and transplanted eelgrass ZosteraMarina meadows
criteria for mitigation. Marine Ecology Progress Series, 65,
251264.
Garcia, A.M., Vieira, J.P., Winemiller, K.O. & Grimm, A.M.
(2004) Comparison of 198283 and 199798 El Nino effects
on the shallow-water fish assemblage of the Patos Lagoon
estuary (Brazil). Estuaries, 27, 905 914.
Gaudreau, N. & Boisclair, D. (2000) Influence of moon phase
on acoustic estimates of the abundance of fish performing
daily horizontal migration in a small oligotrophic lake.
Canadian Journal of Fisheries and Aquatic Sciences, 57,
581590.
Gonzalez, A., Lawton, J.H., Gilbert, F.S., Blackburn, T.M. &
Evans-Freke, I. (1998) Metapopulation dynamics, abun-
dance, and distribution in a microecosystem. Science, 281,
20452047.
Guidetti, P., Fanelli, G., Fraschetti, S., Terlizzi, A. & Boero, F.
(2002) Coastal fish indicate human-induced changes in the
Mediterranean littoral. Marine Environmental Research,
53, 7794.
Hartney, K.B. & Grorud, K.A. (2002) The effect of sea
urchins as biogenic structures on the local abundance of a
temperate reef fish. Oecologia, 131, 506513.
Herben, T., Mandak, B., Bimova, K. & Munzbergova, Z. (2004)
Invasibility and species richness of a community: a neutral
model and a survey of published data. Ecology, 85, 3223
3233.
Hooper, D.U., Chapin, F.S., Ewel, J.J., Hector, A., Inchausti,
P., Lavorel, S., Lawton, J.H., Lodge, D.M., Loreau, M.,
Naeem, S., Schmid, B., Setala, H., Symstad, A.J., Vandermeer, J.
& Wardle, D.A. (2005) Effects of biodiversity on ecosystem
functioning: a consensus of current knowledge. Ecological
Monographs, 75, 335.
Hubbell, S.P. (1997) A unified theory of biogeography and
relative species abundance and its application to tropical
rain forests and coral reefs. Coral Reefs, 16, S9S21.
Hubbell, S.P. (2001) The Unified Theory of Biodiversity and
Biogegraphy. Princeton University Press, Princeton.
Ishitobi, Y., Hiratsuka, J.-i., Kuwabara, H. & Yamamuro, M.
(2000) Comparison of fish fauna in three areas of adjacent
eutrophic estuarine lagoons with different salinities.
Journal of Marine Systems, 26, 171181.
Jensen, A.L. (1997) Limiting similarity for coexistence of lake
herring (Coregonus artedii) and chubs (Coregonus hoyi).
Ecological Modelling, 95, 1115.
Joyeux, J.C. & Ward, A.B. (1998) Constraints on coastal
lagoon fisheries. Advances in Marine Biology, 34, 73199.
Justic, D. (1991) A simple oxygen index for trophic state
description. Marine Pollution Bulletin, 22, 201204.
Karim, M.R., Sekine, M. & Ukita, M. (2003) A model of fish
preference and mortality under hypoxic water in the coastal
environment. Marine Pollution Bulletin, 47, 2529.
Kinzig, A.P., Pacala, S.W. & Tilman, D. (2002) The Func-
tional Consequences of Biodiversity: Empirical Progress
and Theoretical Extensions. Princeton University Press,
Princeton.
Knoppers, B. (1994) Aquatic primary production in coastal
lagoons. Coastal Lagoon Processes (ed. B. Kjerfve),
pp. 243286. Elsevier. Science Publishers, Amsterdam.
Leibold, M.A., Holyoak, M., Mouquet, N., Amarasekare, P.,
Chase, J.M., Hoopes, M.F., Holt, R.D., Shurin, J.B., Law,
R., Tilman, D., Loreau, M. & Gonzalez, A. (2004) The
766 metacommunity concept: a framework for multi-scale
D. Mouillot community ecology. Ecology Letters, 7, 601 613.
Leibold, M.A. & Norberg, J. (2004) Biodiversity in metacom-
munities: plankton as complex adaptive systems? Limnology
and Oceanography, 49, 12781289.
Levin, L.A., Boesch, D.F., Covich, A., Dahm, C., Erseus, C.,
Ewel, K.C., Kneib, R.T., Moldenke, A., Palmer, M.A.,
Snelgrove, P., Strayer, D. & Weslawski, J.M. (2001) The
function of marine critical transition zones and the
importance of sediment biodiversity. Ecosystems, 4, 430
451.
Levin, S.A. & Paine, R.T. (1974) Disturbance, patch formation,
and community structure. Proceedings of the National
Academy of Sciences USA, 71, 27442747.
Loreau, M., Mouquet, N. & Gonzalez, A. (2003) Biodiversity
as spatial insurance in heterogeneous landscapes. Proceedings
of the National Academy of Sciences USA, 100, 12765
12770.
Loreau, M., Naeem, S., Inchausti, P., Bengtsson, J., Grime,
J.P., Hector, A., Hooper, D.U., Huston, M.A., Raffaelli,
D., Schmid, B., Tilman, D. & Wardle, D.A. (2001) Ecology
biodiversity and ecosystem functioning: current knowledge
and future challenges. Science, 294, 804 808.
Ludsin, S.A., Kershner, M.W., Blocksom, K.A., Knight, R.L.
& Stein, R.A. (2001) Life after death in Lake Erie: nutrient
controls drive fish species richness, rehabilitation. Ecological
Applications, 11, 731746.
MacArthur, R. & Levins, R. (1967) Limiting similarity
convergence and divergence of coexisting species. American
Naturalist, 101, 377387.
Mancinelli, G., Costantini, M.L. & Rossi, L. (2002) Cascading
effects of predatory fish exclusion on the detritus-based
food web of a lake littoral zone (Lake Vico, central Italy).
Oecologia, 133, 402 411.
McGill, B.J. (2003) A test of the unified neutral theory of
biodiversity. Nature, 422, 881885.
Mills, M.D., Rader, R.B. & Belk, M.C. (2004) Complex
interactions between native and invasive fish: the simultaneous
effects of multiple negative interactions. Oecologia, 141,
713721.
Mouillot, D., Dumay, O. & Tomasini, J.A. (2007) Limiting
similarity, niche filtering and functional diversity in coastal
lagoon fish communities. Estuarine, Coastal and Shelf Science,
71, 443 456.
Mouillot, D., Gaillard, S., Aliaume, C., Verlaque, M.,
Belsher, T., Troussellier, M. & Chi, T.D. (2005) Ability of
taxonomic diversity indices to discriminate coastal lagoon
environments based on macrophyte communities. Ecological
Indicators, 5, 117.
Mouillot, D., Titeux, A., Migon, C., Sandroni, V., Frodello,
J.P. & Viale, D. (2000) Anthropogenic influences on a
mediterranean Nature Reserve: modelling and forecasting.
Environmental Modeling and Assessment, 5, 185 192.
Mouquet, N. & Loreau, M. (2003) Community patterns in
sourcesink metacommunities. American Naturalist, 162,
544 557.
Nystrom, M., Folke, C. & Moberg, F. (2000) Coral reef
disturbance and resilience in a human-dominated environment.
Trends in Ecology and Evolution, 15, 413 417.
Olden, J.D., Poff, N.L., Douglas, M.R., Douglas, M.E. &
Fausch, K.D. (2004) Ecological and evolutionary conse-
quences of biotic homogenization. Trends in Ecology and
Evolution, 19, 18 24.
2007 British Parker, J.D., Duffy, J.E. & Orth, R.J. (2001) Plant species
diversity and composition: experimental effects on marine
Ecological Society
epifaunal assemblages. Marine Ecology Progress Series,
No claim to original
224, 5567.
French government
Peres-Neto, P.R. (2004) Patterns in the co-occurrence of fish
works, Journal of
species in streams: the role of site suitability, morphology
Applied Ecology, 44, and phylogeny versus species interactions. Oecologia, 140,
760767 352360.
Peterson, C.H., Grabowski, J.H. & Powers, S.P. (2003)
Estimated enhancement of fish production resulting from
restoring oyster reef habitat: quantitative valuation.
Marine Ecology Progress Series, 264, 249264.
Peterson, C.H. & Lipcius, R.N. (2003) Conceptual progress
towards predicting quantitative ecosystem benefits of
ecological restorations. Marine Ecology Progress Series,
264, 297307.
Powers, S.P., Peterson, C.H., Christian, R.R., Sullivan, E.,
Powers, M.J., Bishop, M.J. & Buzzelli, C.P. (2005) Effects of
eutrophication on bottom habitat and prey resources of
demersal fishes. Marine Ecology Progress Series, 302, 233
243.
Ramos Miranda, J., Mouillot, D., Flores Hernandez, D., Sosa
Lopez, A., Do Chi, T. & Ayala Perez, L. (2005) Changes in
four complementary facets of fish diversity in a tropical
coastal lagoon after 18 years: a functional interpretation.
Marine Ecology Progress Series, 304, 113.
Rehage, J.S. & Sih, A. (2004) Dispersal behavior, boldness,
and the link to invasiveness: a comparison of four Gambusia
species. Biological Invasions, 6, 379391.
Roberts, C.M. & Hawkins, J.P. (1999) Extinction risk in the
sea. Trends in Ecology and Evolution, 14, 241246.
Roberts, C.M., McClean, C.J., Veron, J.E.N., Hawkins, J.P.,
Allen, G.R., McAllister, D.E., Mittermeier, C.G., Schueler,
F.W., Spalding, M., Wells, F., Vynne, C. & Werner, T.B.
(2002) Marine biodiversity hotspots and conservation
priorities for tropical reefs. Science, 295, 1280 1284.
Sanvicente-Anorve, L., Flores-Coto, C. & Chiappa-Carrara,
X. (2000) Temporal and spatial scales of ichthyoplankton
distribution in the Southern Gulf of Mexico. Estuarine,
Coastal and Shelf Science, 51, 463 475.
Sfriso, A., Birkemeyer, T. & Ghetti, P.F. (2001) Benthic
macrofauna changes in areas of Venice lagoon populated
by seagrasses or seaweeds. Marine Environmental Research,
52, 323 349.
Sibbing, F.A. & Nagelkerke, L.A.J. (2001) Resource partition-
ing by Lake Tana barbs predicted from fish morphometrics
and prey characteristics. Reviews in Fish Biology and Fisheries,
10, 393 437.
Sogard, S.M. (1989) Colonization of artificial seagrass by
fishes and decapod crustaceans importance of proximity
to natural eelgrass. Journal of Experimental Marine Biology
and Ecology, 133, 1537.
Sosa-Lopez, A., Mouillot, D., Ramos-Miranda, J., Flores-
Hernandez, D. & Do Chi, T. (2007) Fish species richness
decreases with salinity in tropical coastal lagoons. Journal
of Biogeography, 34, 5261.
Stachowicz, J.J., Fried, H., Osman, R.W. & Whitlatch, R.B.
(2002) Biodiversity, invasion resistance, and marine ecosystem
function: reconciling pattern and process. Ecology, 83,
25752590.
Statzner, B., Doledec, S. & Hugueny, B. (2004) Biological trait
composition of European stream invertebrate communities:
assessing the effects of various trait filter types. Ecography,
27, 470 488.
Thomas, C.D., Cameron, A., Green, R.E., Bakkenes, M.,
Beaumont, L.J., Collingham, Y.C., Erasmus, B.F.N., de
Siqueira, M.F., Grainger, A., Hannah, L., Hughes, L.,
Huntley, B., van Jaarsveld, A.S., Midgley, G.F., Miles, L.,
Ortega-Huerta, M.A., Peterson, A.T., Phillips, O.L. &
Williams, S.E. (2004) Extinction risk from climate change.
Nature, 427, 145148.
Turpie, J.K., Beckley, L.E. & Katua, S.M. (2000) Biogeography
and the selection of priority areas for conservation of South
African coastal fishes. Biological Conservation, 92, 59
72.
Volkov, I., Banavar, J.R., Hubbell, S.P. & Maritan, A. (2003)
Neutral theory and relative species abundance in ecology.
Nature, 424, 10351037.
Wilson, D.S. (1992) Complex interactions in metacommunities,
767 with implications for biodiversity and higher levels of
Biodiversity selection. Ecology, 73, 1984 2000.
Wilson, J.B. (1999) Guilds, functional types and ecological
management in
groups. Oikos, 86, 507522.
coastal lagoon fish Ziv, Y. (1998) The effect of habitat heterogeneity on species
communities diversity patterns: a community-level approach using an
object-oriented landscape simulation model (SHALOM).
Ecological Modelling, 111, 135 170.
Zobel, M. (1997) The relative role of species pools in deter-
mining plant species richness. An alternative explanation of
species coexistence? Trends in Ecology and Evolution, 12,
266269.
Received 20 January 2006; final copy received 20 March 2007
Editor: Robert Freckleton

2007 British
Ecological Society
No claim to original
French government
works, Journal of
Applied Ecology, 44,
760767

View publication stats