Effect of nursery habitat on density-dependent habitat selection in juvenile flatfish
Ariel L. Camp
a,
⁎, Clifford H. Ryer
b
, Benjamin Laurel
b
, Kelly Seals
c
a
Dept. Biology, Hofstra University, Hempstead, NY 11549, USA
b
Fisheries Behavioral Ecology Program, Alaska Fisheries Science Center, NOAA Fisheries, Hatfield Marine Science Center, Newport, OR 97365, USA
c
The Maritime Aquarium at Norwalk, Norwalk, CT 06854, USA
abstract article info
Article history:
Received 2 August 2010
Received in revised form 2 May 2011
Accepted 3 May 2011
Available online 2 June 2011
Keywords:
Carrying capacity
English sole
Estuary
Northern rock sole
Predation risk
Nursery habitats contribute significantly to adult recruitment of marine fish, but the density of juveniles that
they can support depends in part on conspecific interactions among juveniles. Density-dependent habitat
selection (DDHS) was compared in the laboratory using two species of age-0 yr flatfish: 1) English sole
Pleuronectes vetulus, which utilizes estuarine nurseries at high densities and 2) northern rock sole Lepidopsetta
polyxystra, which occurs at lower densities in coastal nurseries and is known to exhibit DDHS. The pattern of
DDHS was hypothesized to be similar in both species, but to initiate at higher densities and be less responsive
to predation risk in English sole. Trials were conducted over eight density treatments (0.4, 0.8, 1.5, 3.0, 6.1,
12.2, 18.6, and 23.8 fish m
-2
), both with and without predation risk, and offered fish equal access to a
preferred sand habitat and a less-preferred gravel habitat. Rock sole showed typical DDHS by increasing
percentage use of the gravel habitat as conspecific density increased, with DDHS initiated at about
1.5 fish m
-2
. English sole showed no DDHS: there was no relationship between percentage use of gravel
habitat and conspecific density. Predation risk caused rock sole to significantly decrease gravel usage across all
densities, but did not significantly change habitat selection for English sole. These results suggest that each
species may be behaviorally adapted to the conspecific density and predation risk of their respective nursery
habitats, and correspond well with the comparatively risk-prone behaviors of English sole. The relationship
between nursery habitat and density-dependent behavior may be further understood by studies integrating
factors such as turbidity, food resources and ontogeny.
© 2011 Elsevier B.V. All rights reserved.
1. Introduction
Many marine fish species rely on nurseries: juvenile habitats that
contribute a disproportionately large number of recruits to adult
populations by increasing juvenile growth, survival, or successful
migration to the adult habitat (Beck et al., 2001). The importance of
nursery habitats is well studied (e.g., Gillanders, 1997; Krygier and
Pearcy, 1986; Rooper et al., 2004; van der Veer et al., 2000), but the
factors that determine how species exploit nurseries are diverse and
often unquantified. The ability to support comparatively high
densities of juveniles is often considered a hallmark of nurseries
(e.g. Orth and van Montfrans, 1987), but carrying capacity will depend
on the resources available to juveniles, as well as their survival and
density-dependent interactions (Norcross et al., 1995; Rangeley and
Kramer, 1998; Rooper et al., 2003).
Following ideal free distribution theory, juveniles should distrib-
ute themselves between preferred (high quality) and less-preferred
(low quality) habitats such that fitness is comparable between
habitats (Fretwell and Lucas, 1970). Individuals are expected to
move to a less-preferred habitat as the quality of preferred habitat is
degraded by increased conspecific density. Such behavior is defined as
negative density-dependent habitat selection (DDHS). Juvenile
flatfish are a useful model system for studying DDHS behavior:
pelagic juveniles settle into nurseries at comparatively high densities
(Rooper et al., 2003), and the two-dimensional nature of flatfishes'
habitat may increase lateral concentration and density-dependent
forces (Beverton, 1995).
Flatfish habitat preference depends upon sediment type, depth
and temperature (Gibson, 1994; Rooper et al., 2003). Northern rock
sole Lepidopsetta polyxystra (Orr and Matarese, 2000), hereafter ‘rock
sole’, age-0 yr juveniles are most abundant at the shallow depths and
higher temperatures of coastal embayments (Stoner et al., 2007), and
prefer sand or silts (Stoner and Ottmar, 2003). In contrast, age-0 yr
English sole Pleuronectes vetulus (Girard, 1854) are most abundant in
estuaries (Krygier and Pearcy, 1986), with an estimated 50% of adults
on the central California coast recruiting from estuaries, which
comprise only 6% of the available juvenile habitat (Brown, 2006).
No sediment preference data is available for English sole.
Conspecific densities of each species differ between nursery
habitats: rock sole have been observed at 0.25–0.60 fish m
-2
(Hurst
et al., 2007; Lemke and Ryer, 2006a; Stoner et al., 2007) in coastal
Journal of Experimental Marine Biology and Ecology 404 (2011) 15–20
⁎ Corresponding author at: Dept. of Ecology and Evolution, Brown University, 80
Waterman St., Box G-W, Providence, RI 02912, USA. Tel.: +1 401 863 9578; fax: +1
401 863 7544.
E-mail address: ariel_camp@brown.edu (A.L. Camp).
0022-0981/$ – see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.jembe.2011.05.001
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