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SUMMARY: Indicators of growth and condition were used to compare the habitat quality of nurseries of juvenile sole (Solea
solea L.) in the Bay of Biscay, based on one survey in 2000. The four biological indicators are poorly correlated with each other,
suggesting that no single measure may give an adequate description of fish health and of its habitats quality. Growth indicators
showed significant differences between northern and southern areas. Juveniles from the two southernmost nurseries, the Gironde
estuary and the Pertuis Antioche, displayed significant lower otolith increment widths and mean sizes. These differences were
inversely related to water temperature and unrelated to genetic or age differences, and are unlikely to be due to limiting trophic
conditions in the nurseries. Hence, they may be considered in terms of differences in habitat quality and potential anthropogenic
impacts. Condition indices do not show this north-south pattern but highlight low condition values in the Pertuis Antioche. Short-
term and fluctuating biochemical indicators such as RNA/DNA ratios appeared to be unreliable over a long-term study, while
morphometric indices seemed to be relevant, complementary indicators as they integrate the whole juvenile life-history of sole in
the nurseries. The growth and condition indices of juveniles in September 2000 from nursery grounds exposed to the Erika oil
spill in December 1999 were relatively high. These results lead us to suggest that there was no obvious impact of this event on
the health of juvenile sole and on the quality of the exposed nursery grounds a few months after the event.
Keywords: essential fish habitat, otolith daily increment width, RNA/DNA, Erika oil spill.
RESUMEN: CRECIMIENTO Y CONDICIN DE JUVENILES DE LENGUADO (SOLEA SOLEA L.) COMO INDICADORES DE CALIDAD DE HBI-
TAT EN REAS DE CRA COSTERAS Y ESTURICAS DEL GOLFO DE VIZCAYA CON NFASIS EN ZONAS EXPUESTAS AL VERTIDO DEL ERIKA.
- Se usaron indicadores de crecimiento y condicin de lenguados juveniles (Solea solea L.) para comparar la calidad del hbi-
tat de reas de cra en el Golfo de Vizcaya basados en un muestreo de 2000. Los cuatro indicadores biolgicos mostraron bajas
correlaciones entre s, lo que sugiere que no existe una medida nica para describir adecuadamente el estado de los peces y la
calidad de su hbitat. Los indicadores de crecimiento mostraron diferencias significativas entre las reas septentrionales y meri-
dionales. Los juveniles de las reas situadas ms al sur: el estuario del Garona y Pertuis Antioche mostraron incrementos de oto-
litos y tamaos medios significativamente inferiores. Estas diferencias estn inversamente relacionadas con la temperatura del
agua, pero no con diferencias genticas o de edad, y no es probable que se deba a una condicin trfica limitante en las reas
de cra. Por tanto, pueden considerarse en trminos de calidad del hbitat y el potencial impacto antropognico. Los ndices de
condicin no muestras este patrn norte-sur sino que destacan los bajos valores de condicin en Pertuis Antioche. Los indica-
dores bioqumicos variables de corto trmino como las relaciones RNA/DNA no parecieron fiables para estudios de mayor esca-
la, mientras que los ndices morfomtricos parecen ser relevantes y complementarios ya que integran toda el perodo juvenil de
la vida de los lenguados en sus reas de cra. Los valores de crecimiento y condicin de juveniles en septiembre de 2000 en
reas de cra expuestas al vertido de crudo del Erika en diciembre de 1999 fueron relativamente elevadas. Estos resultados sugie-
ren que no hubo un impacto obvio de este evento sobre la salud de los juveniles de lenguado ni en la calidad de las reas de cra
expuestas transcurridos unos pocos meses despus del vertido.
Palabras clave: hbitat esencial de peces, incremento diario de otolitos, RNA/DNA, vertido del Erik.
sm70s1183-15 29/5/06 16:31 Pgina 184
5 56 W
(Dorel et al., 1991). Moreover, the problems of
48
15 selectivity and size-dependent catchability for trawl-
N
site 1 = Bay of Vilaine ingboth very important factors in early summer
site 2 = Loire Estuary are limited at the end of the growing period (Riou et
site 3 = Bay of Bourgneuf
al., 2001).
Samples were obtained with a 3 m beam trawl
site 4 = Pertuis Breton (stretched mesh size of 20 mm) at depths of 5 to 25
Bay of Biscay m at each site. The 3 northern nursery grounds (Bay
site 5 = Pertuis Antioche
of Vilaine, Loire estuary and Bay of Bourgneuf)
were exposed to the Erika oil spill in December
site 6 = Gironde Estuary
1999. All the sole caught were counted and meas-
ured to the nearest cm (total length, TL). The identi-
fication of the 0-group sole was performed using
length-frequency analysis and confirmed from
43
15
examination of otoliths. A total number of 3882 0-
N group juvenile soles were caught during the survey.
08W For each site and for the 0-group sole, 5 individ-
FIG. 1. Localisation of the 6 sampled sites in the Bay of Biscay. uals for each cm length-class in the range of catches
were immediately frozen at 20C. In the laborato-
and estuarine sectors provide highly productive ry, a subsample of 193 0-group juvenile soles was
areas where ichthyofauna is dominated by juvenile used for the growth and condition analyses.
fish of commercial species such as common sole Standard length (SL) was used in the condition
(Solea solea), whiting (Merlangius merlangus), bib analyses to allow for damage to the caudal fin rays.
(Trisopterus luscus), red mullet (Mullus surmeletus) All the survey data are summed up in Table 1.
and wedge sole (Dicologoglossa cuneata), and non-
commercial species such as black goby (Gobius Condition indices
niger), sand goby (Pomatoschistus minutus), drag-
onet (Callionimus lyra) and solennette RNA/DNA ratio
(Buglossidium luteum) (Dsaunay et al. 2003).
Sole juveniles were collected in 6 nursery RNA and DNA concentrations (mg.l-1) were
grounds in the French part of the Bay of Biscay from determined for each individual on a white muscle
the Bay of Vilaine to the Gironde estuary (Fig. 1) sample with the spectrofluorimetric method
during the NURSE survey in September 2000 described by Le Pecq and Paoletti (1966) and mod-
(R.V. Gwendrez). Autumn was chosen because it ified by Karsten and Wollenberger (1977) using
corresponds to the end of the growing season of ethidium bromide and type I DNA purified from calf
juvenile sole in these nursery grounds. At this time, thymus as a standard. For each individual, a sample
the distribution of juvenile soles is representative of of white muscle was homogenised in ice-cold phos-
the productive period and these YoY fish have not phate-buffered saline (PBS) with a Potter and a
moved from the nursery since their settlement Polytron homogeniser. Two 0.5 mL-aliquots were
TABLE 1. Samples description: 0-group juvenile soles from total catches from the survey and for laboratory analyses (RNA/DNA,
condition factor (R), and recent growth index (RG)) with their size ranges (Total Length TL (cm)).
used to measure DNA total nucleic acids, which not include the nucleus. To ensure measures of the
meant adding heparin to dissociate the nucleopro- radius, all sagittae were first polished by hand on
teins, plus ethidium bromide, and then measuring one side in the sagittal plane using aluminium pow-
total fluorescence. Two more aliquots were used to der (0,1m) until the mid-plane (including the
measure the DNA concentration by adding 0.5 mL nucleus) was reached. The radius was measured
of heparin plus 0.5 mL of RNAse (purified from under a stereomicroscope (magnification 24 x) con-
bovine pancreas) to digest the RNA (30 minutes nected to an image analysis system. The right sagit-
incubation), leaving only the DNA. A fifth aliquot tae were then polished again past the mid-plane,
was used to measure background fluorescence; no until the peripheral increments were reached. The
ethidium bromide was added to this aliquot. A sixth otoliths were then etched for 3-4 minutes with 0.1 M
blank aliquot containing only PBS was run each day EDTA (ethylenediamine tetraacetate) (5%, pH =
to calibrate the spectrofluorimeter. Fluorescence 7.5) and rinsed with distilled water. Because we
was assessed on a KONTRON SFM-25 fluores- were not able to reach the same number of outer
cence spectrophotometer. A 360 nm filter was used increments for all otoliths, we chose to measure the
for excitation, and emission was set at 590 nm. 10 outer ones. Increment widths were measured 3
times for each otolith as described above using a
Condition factor (R) light microscope (magnification 312.5 x), and the
average was used for the statistical analysis.
Standard fish length (SL) measured to the nearest Because the increment widths of all otoliths were far
mm and wet weight (W) measured to the nearest mg greater than the light microscopy resolution limit of
were recorded. We examined the residuals derived 0.2 m (Campana et al., 1987), SEM techniques
from the logged length-weight relationship, i.e. the were not required.
deviation of predicted weight estimated from the
common logged length-weight relationship in terms Prerequisities of otolith microstructure analysis
of general fish condition (Blackwell et al. 2000). We
computed the condition factor according to R = log Validation of daily increment deposition
W log Wc, where W is the wet weight observed The daily increment deposition of sole sagittal
and Wc is the computed weight according to the otolith was validated on larval stages by Lagardre
logged length-weight relationship. R gives the rela- (1989) and on juvenile stages by Lagardre et al.
tive condition of each fish compared with the aver- (unpublished data).
age. The condition of fish is proportional to the
residual values. Otolith growth fish growth relationship
We found a significant linear relationship
Recent growth index (RG) between the maximum radius of the right sagittal
otolith and the standard length (SL) of the analysed
Otolith preparation and measurements juveniles (Fig. 2). Thus, the otolith growth, in terms
of increment widths, reflects somatic growth of 0-
Sagittal otoliths were used for the microstructure group juvenile soles (Campana and Neilson, 1985).
analysis. Only one sagitta was used because a pre-
liminary examination showed the left and right Daily increment width otolith size relationship
sagittae to be identical with regard to the microstruc- The increment width is closely related to the
ture. We chose the right sagitta because the antero- otolith radius with marginal (border) increments
dorsal/postero-ventral axis provides the maximum tending to be wider. Hovenkamp and Witte (1991)
growth axis due to nucleus off-centering. All incre- advise using the residuals derived from the logged
ment width measurements were made as far as pos- increment width/otolith radius relationship to com-
sible along this axis. The right sagittae were then pare different growth patterns. As explained above,
removed, cleaned in distilled water, and dried in an we standardised all measurements of increment
oven (24 h at 60C). They were mounted with the widths to the ten marginal daily increments. Hence,
convex side up on standard microscope slides and we compared increments formed at approximately
embedded in glue (cyanoacrylate). Because sagittal the same radii for each individual at each site. We
otoliths are convex, the maximal growth axis does did not find any significant linear relationship
140 Y = 0. 091 x - 15.65 RNA/DNA ratio, and 0-group mean length and
R2 = 0.77
120 N = 193 density between the 6 sampled sites. One way
Standard length (mm)
TABLE 2. ANOVA results for comparisons between the 6 sampled sites: RNA/DNA ratios, R condition factor (mg), RG recent growth index
(mean width of the last 10-day increments (m)), L total length (cm) and D density (ind/10000 m2) of total 0-group juvenile sole catches com-
puted from the survey and T bottom temperature (C); d.f. = degrees of freedom; S.S.= sum of squares; F = Fisher score;
p = p-values of the test.
9
ences between sites (Table 3). Juveniles from the
7 Bay of Vilaine (1) had significantly higher sizes val-
5 ues than those from the Loire estuary (2) and the
1 2 3 4 5 6
Pertuis Breton (4), which in turn showed a greater
size than those from the Bay of Bourgneuf (3), the
(d) Pertuis Antioche (5) and the Gironde estuary (6)
15
(Table 2, Fig. 3d). A significant positive linear rela-
tionship was found between increment widths and 0-
L (cm)
10
group mean total length (Table 3).
5
0
1 2 3 4 5 6 DISCUSSION
FIG. 3. Differences between the sites for the indices measured in
193 0-group soles: (a) the biochemical index RNA/DNA, (b) the Use of condition indices
condition factor R (mg), (c) the recent growth index RG as mean
width (m) of the last 10-day otolith increments and for an addi-
tional indicator computed from the total catches during the survey: Biochemical indicator
(d) 0-group soles length L (total length (cm)).
Nucleic acids have provided useful tools for
assessing the instantaneous growth rate and condi-
Condition factor (R) tion of both larval and juvenile fish (Fukuda et al.,
2001; Kuropat at al., 2002). Several authors have
The ANCOVAs showed significant differences suggested that the RNA/DNA ratio is more related
between the sites (Table 2) and multiple compar- to the nutritional condition than to the growth
isons showed that individuals from the Bay of (Miglavs and Jobling, 1989; Bergeron, 1997). In
Vilaine (1) and the Gironde estuary (6) significantly the present study, this index was not correlated
displayed the greatest values for this indicator with the recent growth index or with 0-group mean
(Table 2, Fig. 3b). Weak but significant relationships length. The estimations of the RNA/DNA ratio
(Table 3) were found between the condition factor showed that juvenile soles from the Bay of Vilaine
and the 0-group juveniles mean density (negative and the Pertuis Antioche displayed lower values for
relationship), and between the condition factor and this index than those from the other nursery areas.
the bottom temperature (positive relationship) Nevertheless, the important result was that for each
(Table 3). However, the condition factor was not site, the mean RNA/DNA ratio was greater than 2,
correlated with the other indicators (Table 3). which is supposed to be a critical threshold of food
TABLE 3. Correlation coefficients between RNA/DNA ratios, R condition factor (mg), RG recent growth index (mean width of the last 10-
day increments (m)), L total length (cm) and D density (ind/10000 m2) of total 0-group juvenile sole catches computed from the survey, T
bottom temperature (C) and latitude.
RNA/DNA R RG L D T latitude
RNA/DNA - - - - - - -
R - - - 0.033* 0.032* -
RG - 0.175** - 0.177** 0.283***
L - - 0.210*** 0.627***
D - 0.378*** 0.131***
T - 0.667***
deprivation for juvenile sole (Richard et al., 1991). Though the highest densities of 0-group juvenile
This suggested that juvenile soles find favourable soles were found in the nursery area located in the
feeding conditions in each nursery ground in Pertuis Breton, there is only a low correlation
autumn, at least in the short term. Comparable between these two parameters. Our results did not
results were obtained in the southern Bight of the provide evidence for density-dependent effects on
North Sea by Gilliers et al. (2004) using the same juvenile sole condition.
indicator. Studies on juvenile soles generally con-
clude that growth rates of the 0-group in nurseries Use of growth indicators
are generally maximal and that food is present in
excess and therefore never growth-limiting Daily patterns recorded in otolith microstructure
(Rogers, 1994; Amara et al., 2001). The same was are useful for measuring growth rates of fish
concluded for the growth of 0-group plaice in dif- (Campana and Neilson, 1985). Several studies have
ferent nursery grounds (e.g. Pihl et al., 2000; demonstrated that it is possible to estimate the
Amara and Paul, 2003). Nevertheless, the growth rate by measuring increment widths
RNA/DNA ratio is a short-term indicator because (Hovenkamp and Witte, 1991; Stunz et al., 2002).
its value can vary from 30 to 40 h after feeding (J.P. Key assumptions for these uses are that the distance
Bergeron, pers. comm.). High frequency assess- between increments is proportional to the somatic
ments of the RNA/DNA ratio would be required to growth rate and that increments occur daily
provide a reliable estimation of the juvenile nutri- (Campana and Neilson, 1985), which were verified
tional status over a study lasting several months. In for this study. Spatial comparisons of the recent
this type of study, it may be more appropriate to growth index measured on juvenile soles in the Bay
use storage indices such as the triacylglycerol/ of Biscay revealed consistent variations in growth
sterol ratio (Fraser, 1989; Suthers et al., 1992; between nurseries. Individuals from the two south-
Lloret et al., 2003), which may be more integrative ernmost nurseries areas, the Pertuis Antioche and
of the past feeding history. the Gironde estuary, showed narrower increment
widths and are therefore likely to have slower
Condition factor (R) growth rates than individuals from northern areas. 0-
group mean length appeared to confirm this result as
The study of condition is a common practice in individuals from these two sites are smaller.
fisheries ecology and is usually based on length- However, it was impossible to get the absolute age
weight data. Morphometric parameters provide sim- of juveniles from otolith microstructure analysis
ple condition indices which are still widely used because of its shape. Differences in common sole
(Suthers, 1998). The key assumption is that for a spawning periods in the Bay of Biscay have been
given length, a heavier fish is in better condition. We reported by Arbault et al. (1986). However, the spa-
used the residuals derived from juveniles logged tial and temporal distributions of sole eggs did not
length-weight relationship as a condition factor. appear to display a latitudinal pattern. Hence, the
Regardless of fish size, it should be a reliable index gradient of growth may not be age-dependent. Both
(Suthers, 1998). The condition of juveniles varied genetic and environmental factors are known to
between the 6 nursery areas but did not show a con- affect the rate of somatic growth in fish. Previous
sistent pattern. Juveniles from the Pertuis Antioche studies (Autem and Kotulas, 1987; Exadactylos et
displayed the lowest values for this index. al., 2001) showed that the juvenile soles caught in
different nurseries in the Bay of Biscay were genet- (McCormick and Molony, 1993; Suthers et al.,
ically homogeneous, which suggests that 0-group 1992; Suthers 1998). Therefore, the choice of the
juvenile growth differences between nurseries may indicators needs to be discussed and depends on the
be due neither to genetic heterogeneities nor to dif- type of study. For example, in the present study,
ferences of temperatures between sites (Le Pape et short-term and fluctuating biochemical indices such
al., 2003). Hence, this geographical gradient in bio- as the RNA/DNA ratio do not appear to be useful.
logical performance is not explained by known nat- On the other hand, morphometric indices based on
ural factors, and the influence of other factors relat- length-weight data seem to be more relevant indica-
ed to human activities must be considered. tors of fish condition and habitat quality, as they
Concerning the influence of temperature, as in most integrate the whole juvenile life-history in the nurs-
fish species in temperate regions, the growth rate of ery ground. The results obtained from growth indi-
young soles (Solea solea) is related to the water tem- cators at the scale of the year 2000 need to be con-
perature (Fonds, 1975). Higher temperatures are firmed by year-to year comparisons and environ-
more favourable for juvenile sole that inhabit warm mental data related to anthropogenic factors.
shallow waters. Therefore, our results showed that
the recent growth and the length of the juvenile Consequences of the Erika oil spill
were, surprisingly, inversely related to the water
temperature. The oil spill of the Erika occurred in December
To conclude, given our results and the fact that 1999 and involved the spread of 20,000 tonnes of
temperature and food are the two most important fuel over more than 400 km of coast in the French
factors governing fish growth (Fonds, 1975), it part of the Bay of Biscay, in particular the 3 north-
seems unlikely that the low growth indicators found ern nursery areas investigated 9 months later in this
for juvenile sole collected in the nursery areas of the study. The fuel arrivals took different shapes (balls,
Pertuis Antioche and the Gironde estuary resulted patties, layers, etc.) and touched all the environ-
from differences in water temperature, food avail- ments. Data on shellfish collected near the Loire
ability, genetic differences or age differences. estuary area by the Rseau National dObservation
Hence, the low growth performances of juvenile de la Qualit du Milieu Marin (RNO), IFREMER
sole in the southernmost nurseries is not explained and the French Ministry for Ecology and
by known natural factors, and the influence of other Sustainable Development showed that very high
factors such as those related to human activities values of PAH (3 to 5000 g.kg-1 dry weight of
must be considered. In a another study Gilliers et al. shellfish) were reached for several months after the
(in press) examine these growth indicators in rela- oil spill in coastal shellfish-farming areas (the
tion to the contamination of the coastal water with chronic contamination level of this part of the coast
chemical contaminants, notably heavy metals such was situated at around 150 g.kg-1 dry weight for
as cadmium, which has been shown to be highly dis- PAH). Coastal shellfish farming underwent a long
charged in the Gironde estuary and the Pertuis period of closure. Moreover, further studies on con-
Antioche. taminations levels demonstrated that the contamina-
These initial results do not yet enable us to deter- tion profiles analysed 2 years after the Erika oil spill
mine all the factors responsible for the variations in were still different from the initial state (Jeanneret et
the growth and condition of the juvenile soles in the al., 2002).
Bay of Biscay, or to conclude that the juvenile soles According to these results on contamination lev-
find habitats of better quality in the northern nursery els, we hypothesised that 0-group juveniles might
areas. have found damaged habitats when they settled a
few months after the oil spill on exposed nursery
What is a good indicator of anthropogenic grounds. Subsequently, they might have lower
impacts? growth and condition when sampled a few months
later at the end of their first growing season. The dif-
As observed in this study, several authors have ferent growth and condition indices used here are
found a poor correlation between separate indices, affected by a number of divergent sources of vari-
and subsequently advised the simultaneous use of ability, and it is very difficult to draw a quantitative
various indicators to provide reliable estimations conclusion from this approach. Nevertheless, the
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