Diet of marine fish larvae and juveniles that use rocky intertidal pools at the
Portuguese coast
By M. Dias
1
, A. Silva
2
, H. N. Cabral
1
and C. Vinagre
1
1
Centro de Oceanografia, Faculdade de Ci^ encias, Universidade de Lisboa, Campo Grande, Lisboa, Portugal;
2
CVRM-Centro de
Geosistemas, Departamento Engenharia Civil, Arquitectura e Georecursos, Instituto Superior T ecnico, Universidade de Lisboa,
Lisboa, Portugal
Summary
The use of tidal pools during early ontogeny is likely to
enhance growth, condition and survival chances of the tran-
sient marine fish larvae and juveniles that use them. How-
ever, the diet of such individuals within tidal pools is poorly
known; this knowledge is important to understand why such
high numbers of individuals use these environments in spring
and summer on the Portuguese coast. Transient marine fishes
were sampled monthly over a two-year period in four tidal
pools of a rocky reef on the west Portuguese coast. The diet
composition in the tidal pools of the most abundant marine
fish larvae/juveniles, Diplodus sargus and Atherina spp., were
investigated. Stomach and gut contents of 483 individuals
(354 D. sargus ranging in total length from 9 to 87 mm and
134 Atherina spp. ranging in total length from 10 to 31 mm)
were analyzed and dietary indices estimated for the different
developmental stages. The diet of D. sargus composed mainly
harpacticoid copepods, chironomid larvae, ostracods, chiron-
omid adults and amphipods; whereas the diet of Atherina
spp. was mainly harpacticoid copepods, ostracods and gas-
tropods. Along the ontogenetic development, the proportions
and diversity of food items in the D. sargus diet varied.
Importance of the harpacticoid copepods decreased with
increasing fish size, while that of amphipods and isopods
increased. Larvae and juvenile D. sargus and Atherina spp.
found in rocky reef tidal pools are opportunistic consumers
of a wide range of prey and thus take advantage of the high
prey availability in these habitats.
Introduction
Ontogenetic shifts in habitat use are a common pheno-
menon in marine fishes (Beck et al., 2001; Gillanders et al.,
2003) and occur as an adaptive response to reduce predation,
enhance food availability and reduce competition (Clark,
1974; Cushing, 1975; Lenanton, 1982; Boesch and Turner,
1984; Macpherson, 1998). Several species of fish use nursery
grounds in the larval and juvenile stages, moving to subtidal
waters as adults (Randall, 1955; Norris, 1963; Major, 1978).
The use of tidal pools during early ontogeny is likely to
enhance growth, condition and survival chances of the indi-
viduals that use them (Thompson and Lehner, 1976; Mor-
ing, 1986, 1990; Gibson, 1994; Mahon and Mahon, 1994;
Macpherson, 1998; Strydom, 2008). These habitats are
highly structured, with many crevices, holes and macroalgae
that make them spatially complex. In this way they provide
refuge, and have a high biodiversity and abundance of
potential prey. During spring and summer, the tidal pools
warm during the ebb tide, attaining temperatures more
favorable for fish growth than those of subtidal waters
(Beckley, 1985a; Moring, 1986; Amara and Paul, 2003;
Cunha et al., 2007). Rapid growth enables young fishes to
seek refuge and avoid a major part of the size-selective pre-
dation (Pihl and Van der Veer, 1992; Gibson, 1994; Leggett
and DeBlois, 1994; Van der Veer et al., 1994).
Various coastal habitats are well known to be nurseries for
juvenile marine fish (Clark, 1974; Lenanton, 1982). These
include estuaries (Wallace and Van Der Elst, 1975; Day
et al., 1981; Wallace et al., 1984; Costa and Bruxelas, 1989;
Vinagre et al., 2010), coastal lagoons (Franco et al., 2006;
Abecasis et al., 2009; Ribeiro et al., 2012), salt marshes (Vei-
ga et al., 2006), coral reefs (Nagelkerken et al., 2000), man-
groves (Nagelkerken et al., 2000; Dorenbosch et al., 2004),
surf zone reefs (Berry et al., 1982), seagrass beds (Nagelker-
ken et al., 2000; Dorenbosch et al., 2004; Verweij et al.,
2008) and the sandy beach surf zone (Lasiak, 1981, 1983).
Estuarine nurseries are the best documented to date, with
numerous studies from as early as the larval phase of numer-
ous fish species (e.g. Miskiewicz, 1986; Roper, 1986; Boehlert
and Mundy, 1988; Whitfield, 1994; Harris and Cyrus, 2000).
Rocky tidal pools are probably the least studied nurseries for
marine fish, leaving a considerable research void worldwide.
Several studies were developed for rocky tidal pools that
included transient juvenile fish. However, newer studies are
scarce, which indicates that the scientific community has not
focused on these habitats in recent years. Published works
concern transient fish recruitment in the tidal pools of South
Africa (Beckley, 2000) and New Zealand (Willis and Roberts,
1996); species diversity and seasonal changes in Brazil (Bar-
reiros et al., 2004), India (Tsering et al., 2012) and California
(Moring, 1986); community structure in Colombia (Castell-
anos-Galindo et al., 2005), Australia (Griffiths, 2003a), Azores
(Santos et al., 1994), Japan (Murase, 2013), Barbados
(Mahon and Mahon, 1994) and Brazil (Cunha et al., 2008);
spatial and temporal dynamics in Australia (Griffiths, 2003b)
and Japan (Arakaki and Tokeshi, 2006); spatial and vertical
U.S. Copyright Clearance Centre Code Statement: 0175-8659/2014/3005–970$15.00/0
J. Appl. Ichthyol. 30 (2014), 970–977
© 2014 Blackwell Verlag GmbH
ISSN 0175–8659
Received: October 23, 2013
Accepted: January 20, 2014
doi: 10.1111/jai.12450
Applied Ichthyology
Journal of