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2008 SOUTHEASTERN NATURALIST 7(4):651–664
Comparison of Red Grouper Populations from Campeche
Bank, Mexico and West Florida Shelf, United States
Linda A. Lombardi-Carlson1,*, Mark A. Grace2,
and David E. De Anda Fuentes3
Abstract - Epinephelus morio (Red Grouper) is an important component of commercial
fisheries for the United States and Mexico. Fishery-independent long-line
surveys that utilized standardized sampling designs were conducted along the west
Florida shelf, Florida and Campeche Bank, Mexico (2001–2002). Detecting true differences
in abundance and sizes are difficult, but by using standard collection and
aging methodologies, the level of bias was kept at a minimum. Relative abundance
was higher in the Campeche Bank compared to the west Florida shelf (1.74 and 0.63
Red Grouper/100 hook hr, respectively). Fish from the Campeche Bank were also
significantly smaller at length and weight. Differences in length, weight, and relative
abundance of Red Grouper may be because of historical fishing pressure and management
regulations, or the available carrying capacity of each area for Red Grouper.
These differences have a substantial impact on how an overfished population can
recover and be sustained.
Epinephelus morio (Valenciennes) (Red Grouper) is a shallow-water
teleost in the family Serranidae and is distributed along hard bottom
throughout waters off the southeast United States, the Gulf of Mexico,
and Caribbean Sea. Several studies have examined the life history of the
Red Grouper, including age and growth (Johnson and Collins 1994, Stiles
and Burton 1994), reproduction (Brulé et al. 1999, Fitzhugh et al. 2006),
and diet (Brulé and Rodriguez-Canche 1993). Red Grouper is a protogynous
hermaphrodite with spawning occurring from January through May
in the Gulf of Mexico. Unlike most groupers, Red Grouper do not appear
to spawn in aggregations (Coleman et al. 1996). Existing tag-recapture
data indicate Red Grouper do not undergo long-distance migrations (<40
nautical miles [nm]) and likely exist in sub-populations (i.e., fishing
stocks) throughout the Gulf of Mexico (Moe 1966, 1972).
Red Grouper are a very important commercial fish. Cuban fishers have
harvested Red Grouper since the mid-1800s throughout the entire Gulf of
Mexico (SEDAR 2006a, Tashiro and Coleman 1977), and Red Grouper is
still an important component of commercial fisheries for both the United
1NOAA Fisheries, Southeast Fisheries Science Center, 3500 Delwood Beach Road,
Panama City, FL 32408. 2NOAA Fisheries, Southeast Fisheries Science Center, PO
Drawer 1207, Pascagoula, MS 39568-1207. 3Instituto Nacional de la Pesca, CRIP
Yucalpeten, A.P. 73 Progresso, Yucatan, Mexico C.P. 97320. *Corresponding author
652 Southeastern Naturalist Vol. 7, No. 4
States and Mexico today. This species comprises approximately 80% of
the grouper fishery (Brulé and Déniel 1996, Schirripa et al. 1999). Red
Grouper landings also differ extensively between these two Gulf of Mexico
areas with annual (1986–2000) fish landings in the Campeche Bank 3.5
times higher than landings in the west Florida shelf (Giménez-Hurtado
et al. 2005, NMFS 2002). On an average, 11,000 metric tons (mt) of Red
Grouper are harvested from the Campeche Bank compared to only 3000 mt
from the west Florida shelf (Giménez-Hurtado et al. 2005, NMFS 2002).
Furthermore, the United States and Mexico have differed in fishery regulatory
measures (i.e., catch and size limits) and historical trends of harvest
The first federal Red Grouper regulations within the United States Exclusive
Economic Zone were sanctioned in 1990. The United States
federal regulations established a size limit of 508 mm, recreational bag
limits (5 per person), a commercial quota of 4173 mt, and established
gear-specific (long-line vs vertical line) fishing zones (Gulf of Mexico
Reef Fish Management Plan, Amendment 1; SEDAR 2006b). In 2000, all
shallow-water grouper harvesting was restricted from February 15th–
March 15th to protect spawning fish. Further, efforts to reduce overfishing
and restore the stock facilitated the commencement of a 10-yr rebuilding
plan that reduced shallow-water grouper complex quotas to 3991 mt and
established a separate Red Grouper quota of 2404 mt (NMFS 2004a).
Since the establishment of a separate Red Grouper quota, all commercial
landings of any species classified in the shallow-water grouper complex
was restricted in 2004 and 2005 (NMFS 2004b, 2005). During the most
recent US stock assessment, Red Grouper was classified as not
overfished and not undergoing overfishing (SEDAR 2006b), a slight improvement
from earlier assessments (NMFS 2002).
The Red Grouper fishery of the Campeche Bank was historically harvested
by three fishing fleets: Mexican minor (artisanal), Mexican major
(mid-sized), and Cuban (see Arreguín-Sánchez et al. 1996, Giménez-
Hurtado et al. 2005). There is no official federal management plan for
Red Grouper in Mexico despite it being the target species of these fleets.
Mexican fleets have free access to the fishery, but the Cuban fleet catch
quota was reduced from 10,000 mt in 1976 to 850 mt in 2002. A minimum
size limit of 300 mm was established in 1999 (Giménez-Hurtado et al.
2005), and a seasonal closure (February 15th–March 15th; Burgos-Rosas
et al. 2003) was recently established to protect spawners. In 1999, the
Cuba-Mexico experts committee considered the Red Grouper fishery overexploited
due to decreasing catch-per-unit-effort (CPUE) as documented
from research cruises (Giménez-Hurtado et al. 2005) and commercial logbook
data (Hernandez and Seijo 2003).
Because of interests in Red Grouper stocks by commercial and recreational
fisheries management initiatives, two fishery-independent long-line
2008 L.A. Lombardi-Carlson, M.A. Grace, and D.E. De Anda Fuentes 653
surveys were conducted along the west Florida shelf, FL and the Campeche
Bank, Mexico during summer 2001 and 2002. These standardized surveys
allowed for the direct comparison of the abundance and age structure of
Red Grouper from two areas in the Gulf of Mexico subjected to varying
environmental and fishery conditions. Our objectives were to compare catch
rates and life-history parameters (age and growth) from Red Grouper using
equivalent sampling and aging methodologies between these two Gulf of
Fishery-independent long-line surveys were conducted by the NOAA
R/V OREGON II in the northeastern Gulf of Mexico (west Florida shelf)
and by the Mexican R/V ONJUKU in the southeastern Gulf of Mexico
(Campeche Bank) using standardized sampling designs and gear (Grace et
al. 2004) in the summer (June, September) of 2001 and 2002.
The gear consisted of a monofilament mainline (426 kg, 1 nm length),
two radar refl ector buoys (attached at the start and end of the mainline),
5-kg weights clipped on the mainline at the beginning, mid-line, and end
to enable the mainline to rest on the ocean fl oor, and 100 gangions (3.66 m
length) with #15/0 circle hooks clipped to the mainline at a spacing of one
per tenth of a nautical mile. Gangions were baited with Scomber scombrus
L. (Atlantic Mackerel; approximately 0.12 kg per hook) in the west Florida
shelf and with Sarda sarda (Bloch) (Atlantic Bonito; approximately 0.12 kg
per hook) in the Campeche Bank. These are the baits of choice of the commercial
fishers in the respective areas. As gear was deployed, vessel speed
ranged from 5 to 7 knots; the gear was allowed to soak for one hour. Vessel
speed was approximately 4 knots during gear retrieval.
Sampling sites were randomly selected among three depth strata (9–55 m,
56–183 m, and 184–366 m; Grace et al. 2004). The number of sites randomly
selected within 60 nm of contiguous sampling zones was proportional to the
area of continental shelf within each depth stratum. Based on estimates of
shelf area within the strata, 50% of sites were allocated in the first depth stratum,
40% in the second depth stratum, and 10% in the third depth stratum.
However, no Red Grouper were caught in waters >40 m, thus comparisons
were limited to the first depth stratum (9–55 m). Depths were then binned in
10-m increments (e.g., 1–10 m, 11–20 m, 21–30 m, 31–40 m). Capture location
(latitude, longitude) and depth were recorded as the first refl ector buoy
was deployed and retrieved.
Catch-per-unit-effort (CPUE; number of Red Grouper per 100 hook hr)
was calculated by area (west Florida shelf, Campeche Bank) and compared
by depth bin. Log transformed CPUE data were statistically compared between
areas and between areas with depth interactions using a generalized
linear model (R function GLM; R Developing Core Team 2007).
654 Southeastern Naturalist Vol. 7, No. 4
Red Grouper were weighed (kg) and measured (mm fork length [FL],
and total length [TL]), and both sagittal otoliths were extracted. Standardized
aging methodologies were utilized for all age determinations
following Johnson and Collins (1994) and Lombardi-Carlson et al. (2008).
Two readers participated in aging otoliths. The date of capture, reader count,
and edge type were used to assign annual ages (Lombardi-Carlson et al.
2008). Several methods were used to determine the precision among initial
age estimates: (1) average percent error (APE), (2) coefficient of variation
(CV), and (3) percent of readings in agreement (PRA) within ±1 or ± 2 bands
(Campana 2001). The primary reader’s ages were used in further analysis.
Differences in age, total length, and whole weight were examined using
single-factor, random-effects analysis of variance by area (ANOVA;
Zar 1999). Size-at-age data were also compared between areas using
ANOVA; comparisons were restricted to age classes with at least five
samples per area.
The R/V OREGON II made 157 long-line sets on the west Florida shelf
(30o–25oN, 86o–82oW; Fig. 1) during September of 2001 and 2002. The R/V
ONJUKU made 68 long-line sets on the Campeche Bank (22o–19oN, 91o–
90oW; Fig. 1) during June and July of 2001 and 2002.
Figure 1. Map of the Gulf of Mexico showing sampling locations from the west
Florida shelf and the Campeche Bank.
2008 L.A. Lombardi-Carlson, M.A. Grace, and D.E. De Anda Fuentes 655
Relative abundance (CPUE) of Red Grouper differed by area and depth.
Surveys on the west Florida shelf captured 99 Red Grouper in 157 sets
(CPUE = 0.63 grouper per 100 hook hr) while surveys on the Campeche
Bank collected 118 Red Grouper in 68 sets (CPUE = 1.74 grouper per 100
hook hr). Regardless of area, the highest CPUE occurred in the 11–20 m
depth bin (1.32 and 3.41 west Florida shelf and Campeche Bank, respectively;
Fig. 2). Red Grouper abundance was significantly different between
areas (GLM: df = 224, t value = -4.43, P < 0.001) and between areas with
depth interactions (GLM: 1–10 m: df = 28, t value = 0.79, P = 0.43; 11–20 m:
d.f. = 68, t value = -3.40, P < 0.001; 21–30 m: df = 47, t value = -1.72, P =
0.24; 31–40 m: d.f. = 18, t value = -7.83, P < 0.001).
The size ranges of Red Grouper caught were similar between areas (285
mm–880 mm TL), but significant differences were found in average size
(Fig. 3). Red Grouper from the Campeche Bank were significantly shorter
(df = 211, F = 10.94, P < 0.001; mean ± s.d.: 480 ± 110 mm TL at Campeche
Bank versus 532 ± 123 mm TL at west Florida shelf) and smaller in weight
(1.6 ± 1.4 kg at Campeche Bank versus 2.5 ± 2.0 kg at west Florida shelf)
than fish collected on the west Florida shelf (ANOVA df = 210, F = 12.71,
P < 0.001).
Red Grouper were aged (n = 151) using otoliths. The average percent
error (APE) between readers for all samples was 5.3%, the coefficient of
variation (CV) was 7.5, and the percentage of readings in agreement (PRA)
Figure 2. CPUE (number of Red Grouper/100 hook hour) for Red Grouper from the
west Florida shelf and the Campeche Bank by depth. Asterisk indicates significant
differences between areas (P < 0.001), as determined from ANOVA.
656 Southeastern Naturalist Vol. 7, No. 4
Figure 3. Red Grouper from the west Florida shelf and the Campeche Bank (a) total
length, (b) whole weight, and (c) age-frequency histograms.
2008 L.A. Lombardi-Carlson, M.A. Grace, and D.E. De Anda Fuentes 657
within ±1 band was 83%, increasing to 95% within ±2 bands. The indexes
of aging error from each area differed greatly (west Florida shelf: APE =
2.0%, CV = 2.9, PRA ± 1 band = 94%, PRA ± 2 bands = 98%; Campeche
Bank: APE = 10.7%, CV = 15.2, PRA ± 1 band = 67%, PRA ± 2 bands =
89%). Annuli (opaque zones) from the Campeche Bank otoliths were faint
with a feathery appearance compared to the clear annuli from the west
Florida shelf otoliths (Fig. 4), which influenced otolith interpretations and
the indexes of precision.
Overall, there was no significant difference in mean age between areas
(ANOVA: df = 150, F = 3.12, P = 0.08; Fig. 3c). Red Grouper from the west
Florida shelf were slightly older than fish from the Campeche Bank (6.4 ± 2
yr versus 5.6 ± 3 yr). Of the age classes (age 5, 6, and 7) with sufficient samples
(n ≥ 5), no significant differences were found in length and weight of
fish at age 6 (length, ANOVA: df = 27, F = 1.51, P = 0.23; weight, ANOVA:
df = 27, F = 0.58, P = 0.46) and at age 7 (length, ANOVA: df = 17, F = 0.06,
P = 0.80; weight, ANOVA: df = 17, F = 1.02, P = 0.33). Age-5 Red Grouper
from the Campeche Bank were significantly smaller in length (ANOVA: df
= 35, F = 7.54, P < 0.01; mean ± s.d.: 488 ± 61 mm TL at Campeche Bank
versus 549 ± 70 mm TL at west Florida shelf) and weight (ANOVA: df = 35,
F = 10.45, P < 0.01; mean ± s.d.: 1.52 ± 0.55 kg at Campeche Bank versus
Figure 4. Red Grouper annuli patterns from whole (13 x, left) and sectioned (25 x,
right) otoliths from (a) the west Florida shelf and (b) the Campeche Bank. Both fish
were age 5 yr.
658 Southeastern Naturalist Vol. 7, No. 4
2.38 ± 0.91 kg at west Florida shelf) compared to age-5 Red Grouper from
the west Florida shelf (Fig. 5).
Figure 5. Size-at-age plots (mean ± s.d.) for (a) length and (b) weight from the west
Florida shelf and the Campeche Bank. Samples sizes greater than five for each area
and age class.
2008 L.A. Lombardi-Carlson, M.A. Grace, and D.E. De Anda Fuentes 659
Our results indicate differences in size and relative abundance of Red
Grouper between the west Florida shelf and Campeche Bank. Fish from the
Campeche Bank were significantly smaller in length and weight. Size-at-age
comparisons were limited because of sample sizes and the quality of increment
clarity; nonetheless, age-5 Red Grouper from the Campeche Bank were
significantly smaller in length and weight. Relative abundance was also
higher in the Campeche Bank compared to the west Florida shelf (1.74 and
0.63 grouper per 100 hook hr, respectively). Detecting real differences in
abundances and size estimates is difficult due to methodological differences
(Cailliet et al. 1990); however, variations due to sampling bias and aging
methodologies were sufficiently controlled, as the same sampling gear and
methods were used in both areas of the Gulf of Mexico.
Unfortunately, our second objective—comparing size-at-age data between
the areas—was not executed. Our efforts to correctly assign annual
ages to fish collected from the Campeche Bank were inhibited by the clarity
of the increments in both whole and sectioned otoliths. Similar problems
in otolith interpretation have been documented for species of pomacentrids
in the tropical western Atlantic (Caldow and Wellington 2003). Otolith
increment clarity has been reported to be affected by the average water
temperature and the annual range in water temperatures characteristic of
subtropical and tropical regions (Caldow and Wellington 2003, Fowler 1995,
Morales-Nin and Panfili 2005). The Campeche Bank (mean ± s.d. = 28 ± 2
oC) average water temperature is 4 oC higher than that of the west Florida
shelf (24 ± 4 oC), and the Campeche Bank has a smaller range in annual
temperatures (Campeche Bank = 23–33 oC, west Florida shelf = 18–33 oC;
NOAA National Data Buoy Center, www.ndbc.noaa.gov). Thus, the higher
average water temperature and lower range in annual water temperature in
the Campeche Bank affected the interpretation of otoliths and the appropriate
assignment to annual age classes, which was refl ected in the indexes of
precision and percent agreement between readers.
The value of comparing catch rates between the two areas contributes
to the understanding of the difference in magnitude of the landings data
(Fig. 6). Because the catch-per-unit-effort data were based on randomized
surveys using standardized methodologies throughout the spatial range of
Red Grouper (Hernandez and Seijo 2003; SEDAR 2006b), the catch rates
can be considered to be in proportion to Red Grouper abundances (Hilborn
and Walters 1992). Therefore, Red Grouper in the Campeche Bank occur
at higher densities but at smaller sizes than Red Grouper along the west
Understanding the factors responsible for the differences in size and
abundance between these two areas is not straightforward. Nonetheless,
these two areas provide the opportunity to evaluate the effects of fishing
660 Southeastern Naturalist Vol. 7, No. 4
on red grouper. As observed from landings data, fishing pressure on the
Campeche Bank is much higher. A classic compensatory response from
intense fishing pressure, as described through density-dependence, is that
as larger adults are removed more readily by fishing gear, population size
will initially decrease, and the available food supply for the smaller fish will
promote an increase in growth rate and therefore a smaller size-at-maturity
(Trippel 1995). Fishers catch the larger, faster growing individuals (Law
2000) and over time, a decrease in fish size and size-at-maturity can occur,
as seen in salmon in the Pacific (Ricker 1981) and Pagrus pagrus (L.) (Red
Porgy) from the US southeastern coast (Harris and McGovern 1997). The effects
of fishing on the Campeche Bank could be responsible for the decrease
in Red Grouper size-at-maturity (509 mm TL [Brulé et al. 1999], 380 mm TL
[Giménez-Hurtado et al. 2003]).
Detecting density-dependence is difficult and requires replicating sampling
in time and space or manipulative experiments that may provide some
evidence of density-dependence (Jenkins et al. 1999, Webster 2003). Our
study was replicated in time and space, but detecting differences between
areas requires a much longer time series since Red Grouper have life spans
up to 30 years (Lombardi-Carlson et al. 2008). Future studies should investigate
density-dependent mechanisms that may explain why differences in
Red Grouper size and abundance exist between these two areas.
Figure 6. Red Grouper landings (combined commercial and recreational) from the
west Florida shelf and the Campeche Bank. Campeche Bank landings data interpreted
from Giménez-Hurtado et al. (2005, Fig. 2). West Florida shelf landings data
from NMFS (2002, Table A1).
2008 L.A. Lombardi-Carlson, M.A. Grace, and D.E. De Anda Fuentes 661
This research was possible through the assistance of the crews of the NOAA R/V
OREGON II and the Mexican R/V ONJUKU and to all the scientific staff, especially
Dean Landi, Lisa Jones, Karen Mitchell, Melissa Cook, and to our Mexican colleague
José Leonardo Castillo-Géniz (Instituto Nacional de la Pesca, CRIP, Ensenada). We
offer our appreciation to John Carlson for his assistance in collecting samples and for
comments on earlier drafts of this manuscript. Finally, our gratitude extends to Marta
Ribera for her knowledge of GIS. Financial support was provided by the US Department
of Commerce. The statements, findings, conclusions, and recommendations are
those of the author(s) and do not necessarily refl ect the views of the Department of
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