954

Abstract.Understanding

the relative importance of pre- and postsettlement processes is critical to understanding the population dynamics of
marine fishes. Our goals in this study
were 1) to examine habitat preference
and habitat use of newly settled Atlantic croaker, Micropogonias undulatus,
and 2) to determine if postsettlement
growth or predation varied with habitat type. Field surveys showed no difference in croaker abundance among
three estuarine habitats: marsh edge,
seagrass, and sand. Behavioral experiments in laboratory mesocosms suggested that the pattern of similar use
of habitats in the field results from a
lack of preference among habitats. In a
field experiment, croaker recruitment
was greater to artificial seagrass than
to sand habitats, but there was no difference in fish density in habitats with
or without food supplementation. Moreover, growth rates were similar in both
sand and artificial seagrass habitats
and in habitats with or without food
supplementation. In a second experiment, we were unable to detect a difference in the density of newly settled
croaker between sand and artificial
seagrass habitats, or between habitats
with predator access limited by cages and
cage controls. Our results demonstrate
that newly settled croaker use different
estuarine habitats similarly, and there
does not appear to be a fitness consequence of using many habitats. We suggest that for habitat generalists, such as
the Atlantic croaker, variability in larval
supply will be a stronger predictor of
population dynamics than will variability of habitat attributes.

Manuscript accepted 21 April 1999.

Fish. Bull. 97:954961 (1999).

Recruitment of Atlantic croaker,

Micropogonias undulatus:
Do postsettlement processes disrupt
or reinforce initial patterns of settlement?*
Rachel Petrik
Phillip S. Levin
Institute of Marine Science
University of California, Santa Cruz
Santa Cruz, California 95064,
E-mail address (for P. S. Levin, contact author): levin@biology.ucsc.edu

Gregory W. Stunz
Department of Marine Biology
Texas A&M University, Galveston, Texas 77553

John Malone
Department of Biology
University of California, Los Angeles, California 90095

Understanding the causes of fluctuations in population abundance is

critical for ecologists and fishery
biologists. For marine fishes with
life histories in which adults have
limited home ranges and larvae are
pelagic and advected vast distances
from natal sites, an understanding of
variability in larval supply to local
populations is critical for understanding the mechanisms that produce dynamics in populations (Caley et al.,
1996). In addition, habitat selection
by settling fish (Carr, 1991; Levin,
1991; Wellington, 1992; Tolimieri,
1995), and habitat-specific growth
and mortality (Heck and Orth, 1980;
Hixon and Beets, 1993; Levin et al.,
1997) may ultimately reinforce or disrupt patterns created by variable larval supply (Jones, 1997). Thus,
knowledge of the degree to which processes such as habitat selection, competition, or predation modify initial
patterns of larval settlement is important in understanding the population dynamics of marine species.
The importance of variability in
postsettlement growth or mortality
and the level to which postsettlement

processes alter initial patterns of

larval settlement can be a function
of habitat structure. For example,
on coral reefs, holes provide a refuge from predation, and on reefs
with large numbers of holes, the
importance of predation is reduced
(Shulman, 1984; Hixon and Beets,
1993). Similarly, Atlantic cod settle
in equivalent densities in a variety
of habitats but suffer lower predation rates in structurally complex
habitats (Tupper and Boutilier,
1995). Thus, habitat-specific mortality disrupts initial patterns of
larval settlement. Differences in
habitat structure may also impact
growth rates or the ability of fish to
procure food (Nelson, 1979; Heck
and Thoman, 1981; Stoner, 1982).
As examples, 1) pinfish have greater
success capturing amphipods in
shoal grass (Halodule wrightii)
than in similar densities of turtle
grass (Thalassia testudinum) (Stoner,
* Contribution 10 of the Partnership for Interdisciplinary Studies of Coastal Oceans
(PISCO): a long-term ecological consortium
funded by the David and Lucile Packard
Foundation.

Petrik et al.: Recruitment of Micropogonias undulatus

1982); 2) Atlantic cod grow faster in seagrass habitats

than in sand, rocky reef, or cobble habitats (Tupper
and Boutilier, 1995); and 3) pinfish exhibit higher
growth rates in seagrass than in sand habitats (Levin
et al., 1997).
Atlantic croaker, Micropogonias undulatus (hereafter referred to as croaker), range from Cape Cod to
Campeche Bank, Mexico (Johnson, 1978), and occur
both offshore and in estuaries in a variety of habitats including mud, sand, and seagrass (White and
Chittenden, 1977; Johnson, 1978; Rooker et al.,
1998). Croaker are an important component of commercial fisheries in the Gulf of Mexico and southeastern United States, often dominating bottom fish
landings, and are an important sport fishery in this
region (Lassuy, 1983). In the Gulf of Mexico, croaker
spawn over the continental shelf or near inlets from
September to May with peak levels occurring before
January (Johnson, 1978; Cowan, 1988; Cowan and
Shaw, 1988). Larval croaker then move toward shore
and may be transported hundreds of kilometers before entering estuarine nursery grounds (Cowan and
Shaw, 1988; Norcross, 1991). In Texas, recruitment of
croaker peaks in November (Rooker et al., 1998). It is
not clear whether variability in abundance of juvenile
croaker is the result of variability in larval supply or
differential postsettlement growth and mortality.
The delivery of larval croaker recruits to estuarine nursery habitats is dependent on large-scale
oceanographic processes (Cowan and Shaw, 1988).
Once fish arrive at estuaries, delivery into suitable
habitats is dependent on currents and tidal processes
(Norcross, 1991). As fish that are competent to settle
approach nursery grounds they have the opportunity to choose specific microhabitats. In this paper
we examined patterns of microhabitat preference and
use by newly settled croaker, as well as the consequences of microhabitat associations. Specifically we
asked 1) Do croaker have specific microhabitat preferences and are these preferences reflected in patterns of abundance in the field? 2) Does food supply
limit the number or growth rates of croaker recruits
in different habitats? 3) Does predation determine
the number of recruits in different habitats?

Methods
Habitat use by newly recruited croaker
To determine what habitats newly recruited Atlantic croaker use, we conducted a field survey during
November 1996 at Christmas Bay (2903'N, 9510'W),
near Galveston, TX. Christmas Bay is a shallow estuary and contains the most easterly well-developed

955

seagrass bed in Texas. A detailed description of this

site can be found in Thomas et al. (1990). The
seagrass bed is dominated by Halodule wrightii with
an average density of 10,469 shoots/m2 (SE=461). An
epibenthic sled was used to quantify fish abundance
in three habitats: bare sand, seagrass meadow, and
marsh edge. We defined marsh edge habitat as the
subtidal substrata directly adjacent to a Spartina
alterniflora marsh. The sled consisted of a 0.66 m
0.5 m opening fitted with a 3-m long net (1-mm mesh)
with a removable codend. Habitats were sampled by
placing the sled on the substratum, extending a 15-m
rope in a semicircular fashion (to avoid disturbing
sampling area) and pulling the sled through a 10-m2
area. Each habitat was sampled four times at two
different sites, resulting in eight samples per habitat type. Differences in croaker density were examined with a two-way analysis of variance with both
site and habitat type as fixed effects. In this and subsequent analyses, if we failed to reject the null hypothesis of no difference in croaker abundance between habitats, then power analysis was performed.
If statistical power was low, we calculated the number of replicates required to achieve sufficient power
to accept the null hypothesis.
To examine habitat preference we performed choice
experiments in laboratory mesocosms. Six 117-L
mesocosms were constructed from round circular
plastic tanks (41.3 cm diameter 60 cm). The
mesocosms were filled with 5 cm of sand, a plastic
mesh screen was placed on top of the sand, and an
additional 5 cm of sand was placed over the mesh.
Each tank was filled with filtered seawater and maintained at ambient light and temperatures. We divided
mesocosms in half, with each half randomly receiving a sand or grass habitat. Sand habitat was the
sand bottom described above. To construct seagrass
habitats, cores of seagrass were randomly collected
from the field and brought to the laboratory where
they were washed and dipped in fresh water. After
leaves were wiped to remove any epiphytic growth,
the cores were planted in each mesocosm.
One croaker (1520 mm SL) was introduced to the
center of each mesocosm and monitored for any abnormal behavior for 24 h. After the initial acclimation period, the location of each croaker was visually
determined hourly for ten consecutive hours. Visual
observations were performed by a single observer
peering into the mesocosm, without disturbing the
fish. This was repeated for six mesocosms over two
days for a total of 12 mesocosm observations. New
fish were used for each trial. Percent occurrence in
each habitat was determined for all twelve trials. A
one-way t-test determined if percent occurrence in
seagrass was different from 50%.

956

Effects of food supply on recruitment and growth

of croaker in varying habitats
Field experiments were conducted in East Lagoon,
located at the eastern most end of Galveston Island,
TX (2920'N, 9444'W). East Lagoon is 1.6 km long,
0.48 km wide, and has a maximum depth of 4.6 m.
Water is exchanged tidally by means of seven 0.92 mdiameter cement culverts with the Galveston Ship
Channel, which runs from the Gulf of Mexico into
Galveston Bay. A detailed description of this site can
be found in Levin et al. (1997). Seagrass, once widespread in Galveston Bay, including East Lagoon, has
decreased by 90% from peak levels (Pulich and White,
1991); no natural seagrass habitats presently are
found in East Lagoon. The absence of natural
seagrass beds allowed us to establish artificial
seagrass beds with desired characteristics, without
the confounding effects of a natural seagrass bed.
Experiments were located >8 m from the Spartina
alterniflora dominated marsh edge and placed at an
average low tide depth of 42 cm.
To test the null hypothesis that food supply does
not limit abundance or growth rates of croaker recruits in different habitats, we conducted an experiment in which food supply was manipulated in sand
and seagrass habitats. On 20 February 1996 we created five blocks each consisting of four 1-m2 experimental plots. Within each block, food supply and
habitat type were manipulated orthogonally. To control for differences in seagrass structure or seagrassassociated food resources, we used artificial seagrass
habitats. Artificial seagrass habitats were constructed from a 1-m2 polyvinylchoride (PVC) (1.3 cm
diameter) frame, strung with monofilament to form
a grid consisting of 576 points. At each of these points
a 16 cm .5 cm strand of green ribbon was woven in,
such that the frame consisted of 576 shoots of
seagrass, each shoot having two leaves. No excessive fouling was observed on the frame or ribbon for
the duration of the experiment. We performed a preliminary experiment to determine if the structure of
the PVC-frame would attract more fish recruits than
bare sand, and no difference was found between the
bare sand plot and the PVC-frame (F2,23=0.512,
=0.61, 1=0.76). Consequently we performed subsequent experiments without a PVC-frame control.
Food supply was experimentally manipulated with
feeding tubes in each experimental plot. Feeding
tubes were constructed of a 7.5-cm diameter length
of PVC pipe attached to a 1.3-cm diameter PVC pipe
stake, with the bottom of the tube about 15 cm from
the substratum, and the top always above the water
line. We provided supplemental food daily for seven
days, from 23 February to 3 March 1996, to half of

Fishery Bulletin 97(4), 1999

the sand and seagrass replicates (i.e. five sand and

five seagrass plots received food). Food consisted of
200 g of fish flesh and 300 mL of water blended to
produce plankton-size particles (Forrester, 1990;
Levin et al., 1997). The fish pure was placed in ice
cube trays and frozen. Each frozen cube yielded
11.8 g of fish flesh. One cube of frozen food was placed
in the feeding tubes of appropriate replicates,
whereas control plots received one ice cube and no
food was added. As the ice cube containing food
melted, it delivered a continuous stream of particles
to the habitat for 515 min. We observed fish readily
consuming the supplemented food in both the field
and laboratory.
On 4 March 1996, the experiment was terminated
by sampling each plot. Recruit density was quantified by using 1 m3 (1 1 1 m) drop samplers
(Zimmerman et al., 1984; Fonseca et al., 1990). Drop
samplers were constructed of 9.5-mm diameter rebar
covered on four sides with taut 2-mm nylon mesh. A
dip net (90 100 cm, 2-mm nylon mesh) was used to
retrieve fish from the samplers, and replicates were
considered adequately sampled when five consecutive passes of the dip net yielded no fish (Fonseca et
al., 1990). A blocked two-factor analysis of variance
was used to test the hypotheses that the abundance
of newly recruited croaker did not vary among habitat or food supplemented treatments.
Five fish from each replicate were haphazardly
selected for further analysis. We measured the 80
selected fish to the nearest 0.1 mm (SL), removed
their otoliths, and stored them in immersion oil for
one week. Fish age was then determined by enumerating the daily growth rings on the lapillar otolith
by using an image analysis system. The existence of
daily rings on croaker otoliths has been validated
previously (Nixon and Jones, 1997). Each otolith was
examined independently three times. If two of the
three counts did not agree, the fish was discarded
and another selected. When two of the three counts
were the same, that count was used as a datum in
the analysis.
Differences in growth rates were examined by using otolith microstructure. Because otolith diameter
was correlated to fish length (r=0.73, n=75), we used
otolith measures as a proxy for growth rate. Measurements were taken inward from the edge of the
otolith to the seventh ring. This distance corresponded to growth during our seven days of food
supplementation. Otolith distances ( ) were then
converted into daily growth rates (mm SL/day), by
using the following equation generated from a regression of otolith diameter on fish length:
Growth = [(otolith distance + 0.002) / 0.014] 7 .

Petrik et al.: Recruitment of Micropogonias undulatus

957

We used a blocked two-factor analysis of variance to

test the hypothesis that growth rates did not vary
among habitat or food supplementation treatments.
Effects of predation on recruitment of croaker to
varying habitats
The null hypothesis that predation has no effect on
recruitment of croaker in sand or seagrass habitats
was tested by using cages to limit predator access to
experimental plots. On 18 March 1996, a fortuitous
seasonal low tide completely exposed the study site
to air, allowing us to erect cages and to ensure that
no recruits or predators occupied the cages at the
start of the experiment. Cages (2 2 1 m) were
constructed with 25-mm mesh on four sides, whereas
cage controls had mesh on two sides. Mesh was large
enough to be transparent to croaker recruits, but fine
enough to prevent predators from entering the cages.
A randomized block design was employed, with habitat (sand or grass) and predator access (cage or cage
control) as fixed effects. Replicates were placed 8 m
apart, and blocks were separated by 10 m. The experiment was terminated after 7 d. Drop samplers were
used to quantify recruits as described previously.

Figure 1
Atlantic croaker densities (mean +1
SE) sampled from three estuarine habitats: marsh edge (edge), seagrass bed
(grass), and barren sand (sand). P values from two-way analysis of variance.

Results
Habitat preference and use by newly recruited
croaker
The density of newly settled croaker (mean TL=14.48,
SE=0.15) did not differ among sand, seagrass, or
marsh edge habitats (F2,18=0.86, =0.44, =0.13)
(Fig. 1). The density of croaker recruits also did not
vary between sites (F1,18=0.09, =0.77) and the interaction between habitat and site was not significant (F2,12=1.07, =0.37).
Our observation on the behavior of croaker recruits
in mesocosms did not reveal a preference between
sand and seagrass habitats (one way t-test, t=1.64,
df=11, =0.13). An average of 36% (SE=16) of the time
was spent in seagrass, and 64% (SE=16) in sand.
Effects of food supply on croaker recruitment and
growth in varying habitats
The abundance of newly settled croaker differed between experimental habitats (F1,11=5.98, =0.03) with
greater recruitment in sand ( X =31.6/m2, SE=6.5)
than in seagrass habitats ( X =13.7/m 2, SE=5.9)
(Fig. 2). Conversely, we did not detect a difference
(F1,11=0.13, =0.73) in the number of croaker in foodsupplemented plots ( X =23.2/ m2, SE=8.3), compared

Figure 2
Atlantic croaker density (mean +1
SE) in 1-m2 artificial seagrass (grass)
and sand habitats, with (food addition) and without (control) food supplementation. P values from a blocked
two-factor analysis of variance.

with control plots ( X =22.1/m2, SE=4.1) (Fig. 2). The

interaction between habitat and food-supplementation was not significant (F1,11=0.96, =0.35). The statistical power of this experiment was low (1=0.18);

958

Fishery Bulletin 97(4), 1999

0.20

0.18

0.16

0.14

0.12

0.1

Figure 3

Figure 4

Growth rate (mean +1 SE) determined from

analysis of otolith microstructure in croaker
from 1-m2 artificial seagrass (grass) and sand
habitats with (food addition) and without (control) food supplementation. P values are from
a blocked two-factor analysis of variance, and
n, the number of fish sampled, is given at the
base of each bar.

Atlantic croaker density (mean +1 SE) in 1 m2

artificial seagrass (grass) and sand habitats
with predators excluded and allowing predatory fish and decapod access (control). P values
from a blocked two-factor analysis of variance.

however, given the small difference in average

croaker density between food-supplementation and
control plots (1.1/m2) and the high within-treatment
variation, we would have needed 55 replicates to
achieve sufficient power (1=0.95) to accept the null
hypothesis of no difference between treatment means.
Growth rates of newly recruited croaker were the
same in sand and grass habitats (F1,70=2.79, =0.09,
<0.001), as well as with or without food supplementation (F1,70=0.26, =0.61, =0.004) (Fig. 3). The interaction between habitat and food-supplementation
on the growth rate of newly recruited croaker was
not significant (F1,70=0.49, =0.49). Growth averaged
0.148 mm SL/day (SE=0.03) in grass and 0.158 mm
(SE=0.03) in sand. Average growth rates of 0.153 mm/
day (SE=0.03) were observed for both control and food
addition treatments (Fig. 3). These growth rates are
similar to growth rates reported elsewhere (Warlen,
1981; Cowan, 1988; Nixon and Jones, 1997), suggesting that our back calculation of growth rates from
otolith measures were not seriously biased.
Effects of predation on recruitment of croaker to
varying habitats
When we examined recruitment of croaker to experimental plots with or without predator access, we were

unable to detect an effect of predation on croaker

recruitment (F1,12=0.01, =0.92) (Fig. 4). Croaker
density averaged 10.7/m2 (SE=3.8) in the caged replicates, and 15.8/m2 (SE=10.1) in cage controls (Fig. 4).
In contrast with the food supplementation experiment, we did not detect a difference in recruitment
between grass and sand habitats (F1,12=0.96, =0.34)
(Fig. 4). The interaction between habitat and predator access was also not significant (F 1,12 =0.10,
=0.76). This experiment also suffered from low
power (1=0.06). Sufficient power to accept the null
hypothesis of no difference in croaker density between cage and cage-control treatments (1=0.95),
would have required 550 replicates.

Discussion
Recruitment of fishes with open populations is affected by variability in larval supply (Jenkins et al.,
1996; Hamer and Jenkins, 1997), habitat selection
by settling larvae (Bell et al., 1987) and postsettlement mortality (Orth et al., 1984), growth (Levin et
al., 1997), and migration (Sogard, 1989). Understanding how these processes interact with each other to
determine population size has been a major focus of
researchers on tropical and temperate reefs (Doherty
and Williams, 1988; Caley et al., 1996) and recently
in seagrass meadows (Bell et al., 1987; Jenkins et
al., 1996; Hamer and Jenkins, 1997). There has also

Petrik et al.: Recruitment of Micropogonias undulatus

been a growing awareness that understanding how

demographic processes vary with habitat structure
will be critical for predicting population size in fishes
that occur in heterogeneous habitats (Levin, 1994).
In this study we examined patterns of abundance of
newly settled Atlantic croaker and demonstrated that
these fish use different estuarine habitats similarly.
The results of our mesocosm experiment suggest that
the pattern of similar recruitment in different habitats results from a lack of preference for specific habitats. In addition, when we investigated habitat differences in postsettlement growth or survivorship,
we were unable to detect strong consequences of using one habitat over another.
We found no evidence suggesting that habitat selection by settling larvae and habitat-specific
postsettlement mortality are important in determining population size in croaker; however, this conclusion is based on nonsignificant statistical tests rather
than explicit acceptance of the null hypothesis of no
difference between treatments. Recent reviews have
stressed the importance of power analysis in detecting a type-II errors (Peterman, 1990; Reed and
Blaustein, 1995; Thomas and Juanes, 1996), and
because we wished to draw conclusions from negative results, power analysis was particularly important. In this study, when a difference between treatment means was not detected, we examined power
in an attempt to determine our ability to accept the
null hypothesis. If the power of the test was too low
to accept the null hypothesis (0.05), the number of
replicates required to achieve this power level was
calculated. For example, no difference was detected
in mean croaker number between plots from which
predators were excluded and control plots, but the
power of this experiment was low (1=0.06). Sufficient power to be able to accept the null hypothesis,
would have required 550 replicates. Our level of replication was inadequate because of the extreme variability in croaker densities among experimental
plotsvariation likely produced by a combination of
stochastic settlement and habitat-specific mortality.
The effect of this variability was to weaken the power
of our experiments to detect small, but real differences among treatments. Although our experimental design precluded the detection of small treatment
effects, the high number of replicates required to
detect these small effects suggests that other processes are likely to be more important in determining variability in abundance.
Although many estuarine species select vegetated
over unvegetated habitat at settlement (Orth et al.,
1984), in some cases, initial patterns of settlement
have little to do with habitat selection by individual
organisms. Settlement may occur at the first suit-

959

able habitat encountered regardless of specific attributes of that habitat (Bell and Westoby, 1986).
Additionally, current patterns may exclude delivery
of competent larvae to some habitats (Morgan et al.,
1996); therefore, even ideal habitats may seldom receive recruits. In such cases, larvae do not select
against a habitat, instead that habitat is never an
available choice. By experimentally providing habitats, and by using a blocked sampling design such
that all habitats were available in a particular location, we eliminated the possibility that settling
croaker would not have the opportunity to choose a
habitat. In our field sampling and experiments,
croaker had the opportunity to choose between vegetated and unvegetated habitats, but they did not
consistently choose one habitat over another. By contrast, in an identical experiment performed at the
same time and in the same study site, pinfish
(Lagodon rhomboides) showed strong responses to
habitat, food supply, and predators (Levin et al.,
1997). Pinfish occurred in much higher densities in
vegetated than in unvegetated habitats and also grew
faster in grass habitats supplemented with food than
in unsupplemented or unvegetated habitats. In addition, the presence of predators reduced pinfish
numbers by 50%. The pinfish and croaker occupying
experimental plots were similar in size (1525 mm
SL), and at this size the diets of the two species are
similar (Darcy, 1985; Soto et al., 1998). Thus, it is
likely that the lack of response by croaker to the habitat attributes we investigated is the result of characteristics of the species rather than an artifact of
sampling or experimental design.
Selection for specific habitats at settlement may
overwhelm variation in larval supply, thus producing variability in recruitment that is associated with
the preferred habitat. This appears to be the case
for pinfish (Levin et al., 1997). Although croaker often form part of fish assemblages within seagrass
(Rooker et al., 1998), they appear to have broad microhabitat preferences, and our results suggest that
there is no strong fitness consequences for croaker
using vegetated versus unvegetated habitats. As a
result, resources associated with the benthic habitat seem unlikely to determine population size of
newly recruiting croaker. Rather, where and when
larvae that are competent to settle are delivered
should determine population size in croaker. The
contrasting results for croaker and pinfish may reflect a more general difference in the processes determining population sizes of fish. For fishes, such
as pinfish, where settling larvae select specific habitats and postsettlement processes reinforce initial
settlement patterns, spatial and temporal variability in habitat should be a strong predictor of future

960

population size. Habitat generalists, such as croaker,

can occupy a range of habitats in any particular location, and thus variability in a specific resource may
not determine population size. Consequently, processes affecting larval supply will be more important
predictors of population size than resource-related
characteristics.
Generalizations about the relative importance of
processes affecting recruitment in fishes have been
elusive (Caley et al., 1996). Although it is widely recognized that understanding the roles of both pre- and
postsettlement is critical, there is still little consensus on the relative importance of different processes
in determining population size. We suggest that neglecting the behavioral and ecological characteristics of individual species may be a major obstacle in
reaching widely accepted generalizations about processes affecting recruitment. By performing similar
experiments on different species we may uncover
generalizations about processes affecting fish populations that have thus far been difficult to attain.

Acknowledgments
We greatly appreciate the assistance in the field provided by T. P. Good and comments on the manuscript
by B. Finley and J. Rooker. Support for this project
was provided by Texas SeaGrant NA56RG0388
project R/F-67 with supplemental support from a
MARFIN grant from NOAA and NSF grant DEB9610353.

Literature cited
Bell, J. D., and M. Westoby.
1986. Abundance of macrofauna in dense seagrass is due
to habitat preference not predation. Oecologia 68:205
209.
Bell, J. D., M. Westoby, and A. S. Steffe.
1987. Fish larvae settling in seagrass: do they discriminate
between beds of different leaf density? J. Exp. Mar. Biol.
Ecol. 111:133144.
Caley, M. J., M. H. Carr, M. A. Hixon, T. P. Hughes,
G. P. Jones, and B. A. Menge.
1996. Recruitment and the local dynamics of open marine
populations. Ann. Rev. Ecol. Syst. 27:477500.
Carr, M. H.
1991. Habitat selection and recruitment of an assemblage
of temperate zone reef fishes. J. Exp. Mar. Biol. Ecol.
146:113137.
Cowan, J. H., Jr.
1988. Age and growth of Atlantic croaker, Micropogonias
undulatus, larvae collected in the coastal waters on the
northern Gulf of Mexico as determined by increments in
saccular otoliths. Bull. Mar. Sci. 42(3):349357.
Cowan, J. H., and R. F. Shaw.
1988. The distribution, abundance, and transport of larval

Fishery Bulletin 97(4), 1999

sciaenids collected during winter and early spring from

the continental shelf waters off west Louisiana. Fish.
Bull. 86(1):129142.
Darcy, G. H.
1985. Synopsis of biological data on the pinfish, Lagodon
rhomboides (Pisces: Sparidae). U.S. Dep. Commer., NOAA
Technical Report NMFS 23, FAO Fisheries Synopsis 141.
Doherty, P. J., and D. McB. Williams.
1988. The replenishment of coral reef fish populations.
Oceanogr. Mar. Biol. Ann. Rev. 26:487551.
Fonseca, M. S., W. J. Kenworthy, D. R. Colby,
K. A. Rittmaster, and G. W. Thayer.
1990. Comparisons of fauna among natural and transplanted eelgrass Zostera marina meadows: criteria for
mitigation. Mar. Ecol. Prog. Ser. 65:251264.
Forrester, G. E.
1990. Factors influencing the juvenile demography of a
coral reef fish. Ecology 71:16661681.
Hamer, P. A., and G. P. Jenkins.
1997. Larval supply and short-term recruitment of a temperate zone demersal fish, the King George whiting,
Sillaginodes punctata Cuvier and Valenciennes, to an
embayment in south-eastern Australia. J. Exp. Mar. Biol.
Ecol. 208(12):197214.
Heck, K. L., Jr., and R. J. Orth.
1980. Structured components of eelgrass (Zostera marina)
meadows in the lower Chesapeake Bay: decapod crustaceans. Estuaries 3:289295.
Heck, K. L., and T. A. Thoman.
1981. Experiments on predator-prey interactions in vegetated habitats. J. Exp. Mar. Biol. Ecol. 53:125134.
Hixon, M. A., and J. P. Beets.
1993. Predation, prey refuges and the structure of coralreef fish assemblages. Ecol. Monogr. 63(1):77101.
Jenkins, G. P., M. J. Wheatley, and A. G. B. Poore.
1996. Spatial variation in recruitment, growth, and feeding of postsettlement King George whiting, Sillaginodes
punctata, associated with seagrass beds of Port Phillip Bay,
Australia. Can. J. Fish. Aquat. Sci. 53:350359.
Johnson, G. D.
1978. Micropogonias undulatus (Linnaeus), Atlantic croaker.
U.S. Fish and Wildlife Service FWS/OBS-78/12, p. 227233.
Jones, G. P.
1997. Relationships between recruitment and postrecruitment processes in lagoonal populations of two coral reef
fishes. J. Exp. Mar. Biol. Ecol. 213:231246.
Lassuy, D. R.
1983. Atlantic croaker. Species profiles: life histories and
environmental requirements (Gulf of Mexico). U.S. Fish
and Wildlife Service FWS/OBS-82-11.3.
Levin, P. S.
1991. Effects of microhabitat on recruitment variation in a
Gulf of Maine reef fish. Mar. Ecol. Prog. Ser. 75(2-3):183
189.
1994. Small-scale recruitment variation in a temperate fish:
the roles of macrophytes and food supply. Environ. Biol.
Fish. 40:271281.
Levin, P., R. Petrik, and J. Malone.
1997. Interactive effects of habitat selection, food supply
and predation on recruitment of an estuarine fish. Oecologia
112:5563.
Morgan, S. G., R. K. Zimmer-Faust, K. L. Heck Jr., and
L. D. Coen.
1996. Population regulation of blues crabs Callinectes
sapidus in the northern Gulf of Mexico: postlarval supply.
Mar. Ecol. Prog. Ser. 133:7388.

Petrik et al.: Recruitment of Micropogonias undulatus

Nelson, W. G.
1979. Experimental studies of selective predation on amphipods: consequences for amphipod distribution and
abundance. J. Exp. Mar. Biol. Ecol. 38:225245.
Nixon, S. W., and C. M. Jones.
1997. Age and growth of larval and juvenile Atlantic
croaker, Micropogonias undulatus, from the Middle Atlantic Bight and estuarine waters of Virginia. Fish. Bull.
95(4):773784.
Norcross, B. L.
1991. Estuarine recruitment mechanisms of larval Atlantic croakers. Tran. Am. Fish. Soc. 120(6):673683.
Orth, R. J., K. L. Heck, and J. vanMonfrans.
1984. Faunal communities in seagrass beds: a review of
the influence of plant structure and prey characteristics
on predator-prey relationships. Estuaries 7(4a): 339350.
Peterman, R. M.
1990. Statistical power analysis can improve fisheries research and management. Can. J. Fish. Aquat. Sci. 47:215.
Pulich, W. M., and W. A. White.
1991. Decline of submerged vegetation in Galveston Bay
system: chronology and relationships to physical processes.
J. Coast. Res. 7: 11251138.
Reed, J. M., and A. R. Blaustein.
1995. Assessment of nondeclining amphibian populations
using power analysis. Cons. Biol. 9(5):12991300.
Rooker, J. R., S. A. Holt, M. A. Soto, and G. J. Holt
1998. Postsettlement patterns of habitat use by sciaenid fishes
in subtropical seagrass meadows. Estuaries 21:318327.
Shulman, M. J.
1984. Resource limitation and recruitment patterns in a coral
reef fish assemblage. J. Exp. Mar. Biol. Ecol. 74:85109.
Sogard, S. M.
1989. Colonization of artificial seagrass by fishes and decapod crustaceans: importance of proximity to natural
seagrass. J. Exp. Mar. Biol. Ecol. 133:1537.
Soto, M. A., G. J. Holt, S. A. Holt, and J. Rooker.
1998. Food habits and dietary overlap of newly settled red

961

drum (Sciaenops ocellatus) and Atlantic croaker (Micropogonias undulatus) from Texas seagrass meadows. Gulf
Research Reports 10:4155.
Stoner, A. W.
1982. The influence of benthic macrophytes on the foraging behavior of pinfish Lagodon rhomboides (L.). J. Exp.
Mar. Biol. Ecol. 58:271284.
Thomas, J. L., R. J. Zimmerman and T. J. Minello.
1990. Abundance patterns of juvenile blue crabs (Callinectes sapidus) in nursery habitats of two Texas bays.
Bull. Mar. Sci. 46(1):115125.
Thomas, L., and F. Juanes.
1996. The importance of statistical power analysis: an example from Animal Behavior. Anim. Behav. 52:856859.
Tolimieri, N.
1995. Effects of microhabitat characteristics on the settlement and recruitment of a coral reef fish at two spatial
scales. Oecologia 102:5263.
Tupper, M., and R. G. Boutilier.
1995. Effects of habitat on settlement, growth, and postsettlement survival of Atlantic cod (Gadus morhua).
Can. J. Fish. Aquat. Sci. 52:18341841.
Warlen, S. M.
1981. Age and growth of larvae and spawning time of Atlantic croaker larvae in North Carolina. Proc. Annu. Conf.
SE Assoc. Fish Wildl. Agencies 34:204214.
Wellington, G. M.
1992. Habitat selection and juvenile persistence control the
distribution of two closely related Caribbean damselfishes.
Oecologia 90:500508.
White, M. L., and M. E. Chittenden Jr.
1977. Age determination, reproduction, and population
dynamics of the Atlantic croaker, Micropogonias undulatus.
Fish. Bull. 75(1):109123.
Zimmerman, R. J., T. J. Minello, and G. Zamora.
1984. Selection of vegetated habitat by brown shrimp
Panaeus aztecus in a Galveston Bay salt marsh. Fish.
Bull. 82: 326336.