Effect of simulated macroalgae on the fish assemblage associated with a temperate
reef system
T. Vega Fernández
a,
⁎, G. D'Anna
a
, F. Badalamenti
a
, A. Pérez-Ruzafa
b
a
CNR — IAMC, Laboratory of Marine Ecology, Via G. da Verrazzano 17, Castellammare del Golfo (TP), 91014, Italy
b
Department of Ecology and Hydrology, University of Murcia, Campus Universitario de Espinardo, Murcia, 30011, Spain
abstract article info
Article history:
Received 2 April 2008
Received in revised form 21 May 2009
Accepted 22 May 2009
Keywords:
Assemblage structure
Artificial algae
Canopy restoration
Habitat complexity
Habitat structure
Repeated measures
Increased habitat complexity is supposed to promote increased diversity, abundance and biomass. This study
tested the effect of the macroalgal cover on temperate reef fishes by mimicking macroalgae on artificial reefs
in NW Sicily (Mediterranean Sea). Macroalgal cover affected reef fishes in different ways and independently
of intrinsic temporal trends. The fish assemblages of manipulated and control artificial reef units differed in
the relative abundances of the associated species, but little in species composition. In line with studies in
seagrass habitats, fishes were most abundant in reefs covered by artificial macroalgae. Three species (Boops
boops, Serranus scriba and Symphodus ocellatus) exhibited consistently greater abundance on vegetated reef
units than on control reef units. The total number of species and the abundance of three particular species
(S. scriba, S. ocellatus and Thalassoma pavo) displayed temporal trends which were independent on short and
large temporal scales. Only fish total biomass and one species (Spicara flexuosa) displayed strong effects of
interaction among the experimental factors. Mechanisms to explain these findings are discussed from
observational evidence on habitat use and interactions among multiple species. This study highlights that
manipulative experiments involving repeated sampling of fish in artificial habitats appear to be a valid
approach to study fish-habitat relationships in fluctuating environments. It is also concluded that macroalgae
mimics may serve as a tool for restoring lost marine vegetated habitats when current human-induced
conditions prevent the recovery of pristine macroalgal stands.
© 2009 Elsevier B.V. All rights reserved.
1. Introduction
Increased habitat complexity has long been recognised to exert a
positive influence on the diversity and abundance of marine
organisms. Habitat complexity promotes resource partitioning by
ameliorating competition (Schoener, 1974), and leads to stabilization
of predator–prey dynamics (Sih, 1987). Structural complexity (sensu
McCoy et al., 1991) is provided by plants in most ecosystems by means
of their density and form (Silvertown, 2004). Thick vegetation is still
permeable for small or appropriately shaped animals whilst larger or
unsuitably shaped species are physically excluded. Hence vegetative
cover is frequently used for refuge and foraging under predation risk
(Lima and Dill, 1990).
In temperate shallow aquatic environments, vascular macrophytes
and macroalgae supply the bulk of the habitat complexity above the
bottom (Ebeling and Hixon, 1991). These marine vegetated commu-
nities support more diverse and abundant fish assemblages than
surrounding unvegetated areas of similar depth, and share many
common species irrespective of vegetation type (Jenkins and
Wheatley, 1998; Guidetti, 2000). This suggests that many fish species
select the structure provided by vegetation rather than specific
vegetative species (Heck et al., 2003). Indeed, field experiments
indicate that plant volume and cover are among the most important
factors driving the structure of temperate fish assemblages (Choat and
Ayling, 1987; Carr, 1989), and are particularly important for small-
bodied fishes (Shulman, 1985; Carr, 1989; Adams et al., 2004; Crawley
et al., 2006).
The effect of plant complexity on fishes may be explained by
habitat related resource availability (Jones, 1991). The type of resource
also exerts great influence on species-specific and size-specific
interactions (Werner et al.,1983b; Persson and Eklöv, 1995). Shelter
and food are the most important niche axes in fishes (Wootton, 1998)
and many species exploit both resources in vegetated habitats
(Lenanton and Caputi, 1989; Crawley et al., 2006). When several
resources are present in multiple habitat types, vegetated areas are
still preferred by some species, thus it is supposed that their individual
fitness is improved there (Werner et al., 1983a; Alofs and Polivka,
2004). More usually, predation refuge and food occur in distinct,
alternate habitat types. The role of plant cover as refuge then takes
precedence in the presence of predators (Werner et al., 1983b;
Shulman, 1985; Gerking, 1994).
How predation success is affected by habitat complexity is strongly
species dependent (Diehl, 1988; Eklöv and Diehl, 1994). In general,
Journal of Experimental Marine Biology and Ecology 376 (2009) 7–16
⁎ Corresponding author. Tel.: +39 092435013; fax: +39 092435084.
E-mail address: tvega@irma.pa.cnr.it (T. Vega Fernández).
0022-0981/$ – see front matter © 2009 Elsevier B.V. All rights reserved.
doi:10.1016/j.jembe.2009.05.012
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