Habitat preferences and site fidelity of juvenile red snapper (Lutjanus campechanus)
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ICES Journal of Marine Science, 59: S43–S50. 2002
doi:10.1006/jmsc.2002.1211, available online at http://www.idealibrary.com on
Habitat preferences and site fidelity of juvenile red snapper
(Lutjanus campechanus)
Ian Workman, Arvind Shah, Dan Foster, and
Bret Hataway
Workman, I., Shah, A., Foster, D., and Hataway, B. 2002. Habitat preferences and
site fidelity of juvenile red snapper (Lutjanus campechanus). – ICES Journal of Marine
Science, 59: S43–S50.
Ten small reefs constructed of either oyster shell or polyethylene webbing and ten
randomly selected open-bottom sites within a distance of 3.7 km of the reefs were used
to determine juvenile red snapper [Lutjanus campechanus (Poey, 1860)] habitat prefer-
ences. The reefs were deployed at 40, 50, 70, and 90 m from a flare stack located in a gas
field off the coast of Mississippi. Juvenile red snapper were observed at one of the
open-bottom sites when it was first surveyed, but none during a second survey con-
ducted 45 days later. Age-0 and age-1 fish showed a preference for the more complex
study reefs, but presence of age-1 appeared to limit recruitment of age-0 to a reef. As
age-1 fish started to leave the reefs, increased numbers of age-0 were observed moving
onto them. However, age-0 were never observed at the 40-m reef occupied by older fish
throughout the study. Distance from the flare stack also appeared to have an effect on
recruitment to the reefs. Age-0 were first observed at the 50-m reefs. They appeared at
the 70-m reefs a week later and at the 90-m reefs almost a month later. Age-1 fish showed
a preference for the reefs located closest to the flare stack. Juvenile red snapper site
fidelity was determined using fish that either were tagged and released on site or were
removed from the capture site before release. Fish from the on-site release were
repeatedly sighted at the capture reef over about a 2-month period. Displaced fish, as
determined with tracking equipment, were able to find their way back to the capture reef
from as far away as 0.43 km in about 25 min. We conclude that juvenile red snapper are
not only faithful to structures, but also have homing capabilities.
Published by Elsevier Science Ltd on behalf of the International Council for the Exploration of
the Sea.
Keywords: habitat preferences, homing instinct, juvenile red snapper, site fidelity.
Accepted 19 November 2001.
I. Workman, D. Foster, and B. Hataway: National Marine Fisheries Service, PO Drawer
1207, Pascagoula, Mississippi 39568-1207, USA; tel: +1 228 762 4591; fax: +1 228 769
8699; e-mail: Ian.Workman@noaa.gov. A. Shah: Merck Research Laboratories, Merck
& Co., Inc., RY33-404, PO Box 2000, Rahway, New Jersey 07065-0900, USA; tel: +1
732 594 8343; fax: +1 732 594 6075; e-mail: arvind–shah@merck.com
Introduction Amendment 9 to the Shrimp Fishery Management
Plan, implemented in May 1998, requires the use of
The red snapper [Lutjanus campechanus (Poey 1860)] by-catch reduction devices (BRDs) in shrimp trawls.
stock in the US Gulf of Mexico (GOM) has been While BRDs have shown good reduction results for
significantly depleted over the past three decades. many species, reduction rates for red snapper are
Contributing to this condition are a combination of marginally acceptable based on requirements established
over-harvesting by commercial and recreational fishers under Amendment 9. There is further concern that
and the high incidental catch of juvenile fish in the GOM trawl-related red snapper mortality may be even greater
shrimp fishery (Schirripa and Legault, 1997). The Gulf than indicated by landings during BRD evaluations.
of Mexico Fisheries Management Council has made Studies on the separation of other species in trawl
some progress in rebuilding the stock by setting com- catches have indicated some mortality resulting from
mercial and recreational fishing quotas and size limits stress and trawl contact (DeAlteris and Reifsteck, 1993;
for adults but has only recently begun to take steps to Broadhurst and Kennelly, 1995; Chopin and Arimoto,
increase juvenile survival. 1995). Net feeding by Atlantic bottlenose dolphin
1054–3139/02/0S0043+08 $35.00/0 Published by Elsevier Science Ltd on behalf of the International Council for the Exploration of the SeaS44 I. Workman et al.
(Tursiops truncatus) and jack crevalle (Caranx hippos) on 3 90 m
by-catch escaping from BRDs has been video-taped by
researchers from the University of Georgia Marine
Reef
Extension Service and the National Marine Fisheries sets 2 70 m
Service (Pascagoula, MS Laboratory). In addition, dis-
placement from bottom structures by trawling activities
1 50 m
could leave the fish without protective habitat and more
likely to be preyed upon. Floated poly
Ensuring increased survival of juvenile red snapper Poly
will require a better understanding of their distribution, Shell
habitat preferences, and behaviour. Most information Double structure
50 m
available has come from trawl studies, and several of
these convey the idea that juveniles are randomly
distributed over the shrimp grounds (Mosely, 1966;
Bradley and Bryan, 1975; Gutherz and Pellegrin, 1988).
m 40
More recent studies show that, from the time they leave Flare stack
the plankton, red snapper are strongly attracted to
structures (Workman and Watson, 1995a, b; Szedlmayer Figure 1. Arrangement of reef types and reef sets relative to the
flare stack adjacent to Chevron MO-861-1.
and Conti, 1999; Bailey, 1995). Their occurrence on
shrimp grounds has been related to the presence of
small structures, including shell, gravel, discarded
materials, burrows in the bottom and squid egg flare stacks and all but two of the primary structures
clusters (Hildebrand, 1954; Workman and Foster, 1994; were removed, causing the study to end.
Szedlmayer and Howe, 1997). Tagging studies were conducted in the gas field and at
We have been studying the occurrence and habits of a small concrete-rubble reef site located 17 km south of
juvenile red snapper on shrimp fishing grounds in the Horn Island Pass (depth about 21 m). This reef (about
northeastern GOM since 1991 (Workman and Foster, 1510 m) was one of a number of small sections of
1994). Workman and Watson (1995a), in the process of rubble that fell adjacent to a larger concrete rubble reef
trying to develop an effective BRD, determined that during deployment.
these fish can be attracted to and concentrated within The comparison between different reef types included
small reefs made of webbing and providing refuge one reef constructed with oyster shell and two con-
(Workman and Watson, 1995b). structed with 40-mm mesh polyethylene webbing using
To learn more about habitat requirements, we com- 13 mm PVC pipe frames (1.51.5 m). The shell reefs
pared the attractive force of different artificial reef types were formed by placing a frame on the bottom and
to that of adjacent open bottom areas. We also studied covering the area within the frame with oyster shell to a
site fidelity to reefs using fish that were tagged and either level approximately 5 cm above the surrounding bot-
released on site or transported from the capture site tom. Both polyethylene reef types were made of webbing
before release. panels attached to the frames. Flexible 15-cm long
polypropylene strands were attached to the webbing
panels at 15-cm intervals to give the effect of seagrass.
One type was floated with a 10-cm diameter plastic float
Materials and methods attached to the centre underside of the webbing; the
other was not floated.
Study sites
Three sets containing one reef of all three types each
Habitat preferences were studied in a gas field (Mobile were deployed along a 360 compass heading at 50, 70,
Block 861) located 12.5 km southeast of Horn Island and 90 m distance from the flare stack in May 1996
Pass off Pascagoula, Mississippi. Initially, the field (Figure 1). The reefs in each set were randomly arranged
contained nine primary structures and five emergency in a row perpendicular to the course heading with
pressure release pipes (flare stacks). The arrangement of in-between distance of 10 m.
structures and submerged pipelines made the area within A third webbing reef type was made of two frames
the field (about 4.3 km2) virtually untrawlable. The stacked one on top of the other and separated by four
study site was located off a flare stack adjacent to one of 15-cm pipe sections installed at the corners. Polyethylene
the primary structures (Chevron MO-861-1) at the west webbing was attached to both frames. The bottom
side of the field. Water depth was about 17 m, and webbing panel was left plain, while the top panel had
bottom type was a mixture of sand and silt over a strands attached as described above and was floated
clay-mud substrate. In September and October 1996, all with a 10-cm diameter plastic float. The double webbingHabitat preferences and site fidelity of juvenile red snapper S45
reef was deployed 40 m away on a 270 heading from the released. One video-taped the experiment (with a Hi 8
flare stack in November 1995. video camera in an underwater housing) from the time
the cage neared the surface to the time the divers either
had lost the fish or had to return to the surface. The
Data collection other diver recorded the time of significant events,
Surveys began in March 1996 on the double webbing monitored course heading with an underwater compass,
reef and in May 1996 on the comparison reefs, and and kept track of the sonic tagged fish with a Dukane
ended in September 1996. A total of 15 double webbing underwater receiver. A surface team operating from a
reef surveys and 11 comparison reef surveys were con- small boat continued to track the fish with a Dukane
ducted at 4-d to 26-d intervals. Visual counts of all red receiver from the point the divers left off. Two follow-up
snapper inhabiting the reefs were made by two divers dives were made on the rubble reef to search for sonic
and recorded on plastic slates. Counts were compared tags with an underwater receiver and to record sightings
on site and, if necessary, repeated until agreement was of fish with colour-identifiable T-bar tags.
reached. Counts were made by year class (distinctly
separated by size) and included fish spawned in 1995
(age-1) and 1996 (age-0). Analysis
To compare the attractive power of the reefs to that of To investigate effects of reef type, reef set, and survey
adjacent areas, ten open bottom sites located within date on age-0 and age-1 red snapper counts, a 3-factor
3.7 km distance were randomly selected and surveyed factorial analysis of variance model (including all two-
using a circle line search method. The line (or radius of way interactions) was fitted. The model tested for each
the circle) was 15 m long, and one end was anchored in year class was:
the middle of the search area. Two divers using the limits
of lateral visibility or 1.5 m increments, whichever was Yijk =+i +j +k +()ij +()ik +()jk +ijk,
shorter, worked their way out as they circled the centre
point to the end of the line. All snapper occurring within where Yijk is red snapper count for reef type (i=1,2,3),
the circle were counted and bottom features were period (j=1,2, . . ., 11), and set no (k=1,2,3), is overall
recorded. mean effect, i is reef type effect, j =period (date) effect,
To determine site fidelity, 21 age-1 red snapper k =set No effect, and ()ij, ()ik, and ()jk are the
inhabiting the double webbing reef were captured in late corresponding two-way interaction effects.
July 1996 (using a 3-m diameter, 2.54-cm mesh, mono- Pair-wise comparisons of the mean counts were made
filament dropnet) and tagged with Floy external T-bar using Fisher’s least significant differences technique. A
anchor tags. They were released on site and monitored multivariate analysis was also performed, but led to the
through re-sightings on reef surveys. In addition, three same conclusions.
horizontal-vertical displacement experiments (simulat-
ing shrimp trawl capture) were conducted in August
1998 with age-1 red snapper captured at the rubble reef Results
site. The same dropnet was used and the snapper caught
were transferred to a 1 m high1 m wide1.5 m long Habitat preferences
collapsible holding cage. Several fish were tagged with Age-1 snapper (64–114 mm FL) were observed at the
T-bar tags (a different colour for each experiment), and two polyethylene reef types from May onwards
during each of the last two experiments a Sonotronics (Table 1). Numbers at the floated reefs peaked in late
sonic tag (2.9 cm long; 1.4 cm diameter) was attached to July and declined markedly in September, while
one fish. The fish were held on site for about 12 h before numbers at the float-less polyethylene reefs fluctuated
displacement. without trend until September, when they disappeared
Just before displacement, divers collapsed the forward completely. Age-1 fish were infrequent visitors at the
end of the cage into a wedge shape and attached two shell reefs.
towing lines. Using underwater propulsion vehicles, the The first age-0 recruit (12.8 mm FL) was observed in
cage was towed between 0.37 and 0.43 km from the mid-July (Table 1) at the 50-m shell reef. Age-0 appeared
capture point, the direction being different for each at the floatless polyethylene reefs 1 wk later and at the
experiment. The first two experiments involved straight- floated reefs almost a month later. Their numbers gradu-
line displacements, the third a dog-leg displacement ally increased through late August and then sharply
pattern with a 45 turn at about the half-way point. At increased at all three reef types. The timing of the
the end of the horizontal track, a float line was attached accelerated increase in age-0 corresponds with the time
and released to the surface. when age-1 started to disappear. Age-0 continued to
A second pair of divers entered the water just before recruit to the reefs through the last survey conducted in
the cage was hauled to the surface and the fish were mid-September.S46 I. Workman et al.
Table 1. Mean red snapper counts by date, reef type (FPR: floated polyethylene reef; PR: polyethylene
reef; SR: shell reef) and age (n=3).
FPR PR SR
Date Age-1 Age-0 Age-1 Age-0 Age-1 Age-0
23 May 1996 6.7 0.0 1.3 0.0 0.0 0.0
18 Jun 1996 14.0 0.0 1.0 0.0 0.0 0.0
03 Jul 1996 12.7 0.0 2.0 0.0 0.0 0.0
19 Jul 1996 16.7 0.0 0.7 0.0 1.0 0.3
26 Jul 1996 23.0 0.0 0.3 0.3 0.0 0.7
05 Aug 1996 15.0 0.0 1.0 2.3 0.0 1.0
14 Aug 1996 13.3 0.3 1.7 3.7 0.0 3.3
27 Aug 1996 5.0 6.7 4.0 13.0 0.3 11.7
04 Sep 1996 11.3 7.3 0.3 14.7 0.0 13.3
11 Sep 1996 1.7 12.0 0.0 12.3 0.0 10.3
19 Sep 1996 1.0 26.0 0.0 39.7 0.0 21.3
Mean (n=33) 10.9 4.8 1.1 7.8 0.1 5.6
Table 2. Analysis of variance for (A) age-1 and (B) age-0 red snapper as dependent variable.
Source DF SS MS F-value p-value
A. Age-1 (R-square: 0.91)
Model 58 4091 71 6.8 0.0001
Reef-type 2 2359 1179 114.3 0.0001
Date 10 460 46 4.5Habitat preferences and site fidelity of juvenile red snapper S47
Table 3. Mean red snapper counts by date, reef set, and age (n=3).
Set 50 m 70 m 90 m
Date Age-1 Age-0 Age-1 Age-0 Age-1 Age-0
23 May 1996 6.0 0.0 1.7 0.0 0.3 0.0
18 Jun 1996 7.3 0.0 3.7 0.0 4.0 0.0
03 Jul 1996 7.0 0.0 3.0 0.0 4.7 0.0
19 Jul 1996 8.0 0.3 6.3 0.0 4.0 0.0
26 Jul 1996 8.0 0.7 7.0 0.3 8.3 0.0
05 Aug 1996 5.3 1.0 6.7 2.3 4.0 0.0
14 Aug 1996 6.0 3.0 5.0 4.0 4.0 0.3
27 Aug 1996 4.0 11.3 3.7 11.7 1.7 8.3
04 Sep 1996 4.3 11.3 6.3 11.7 1.0 12.3
11 Sep 1996 0.0 12.0 0.0 11.7 1.7 11.0
19 Sep 1996 1.0 20.7 0.0 26.7 0.0 39.7
Mean (n=33) 5.2 5.5 3.9 6.2 3.1 6.5
Table 4. Red snapper counts (C), size range (mm), depth (D in m), and bottom features (ss: sand-silt)
at open bottom survey sites (coordinates relative to 30N 88W).
Date Lat. Long. C Range D Features
19 Jul 1996 07.5 27 0 — 14.3 ss
24 Jul 1996 08 28.5 0 — 13.7 ss
24 Jul 1996 07.5 28 0 — 13.7 ss
26 Jul 1096 06 26 0 — 16.8 ss
30 Jul 1996 06.5 25.5 0 — 16.5 ss
05 Aug 1996 07.5 25.5 0 — 16.5 ss with some shells
05 Aug 1996 08 26.5 14 2.8–19.0 14.6 ss with numerous shells and burrows
14 Aug 1996 07.5 25 0 — 18.0 ss with some shells and burrows
11 Sep 1996 06.5 26 0 — 17.4 ss
11 Sep 1996 05.5 25 0 — 18.0 ss
19 Sep 1996 08 26.5 0 — 14.6 ss with some shells and burrows
Table 5. Number of tagged red snapper sighted by date at double webbing reef and flare stack (21 fish
tagged; two detached tags found on 26 July).
Date 26 Jul 30 Jul 05 Aug 14 Aug 04 Sep 11 Sep 19 Sep
Double webbing reef 6 5 5 10 3 3 1
Flare stack 0 0 1 1 1 2 1
and characterized by numerous small shells and mantis during the last survey in mid-September (Table 5). In
shrimp burrows, was surveyed once more in mid- addition, one of the larger tagged fish recruited to the
September, but no red snapper and fewer shells and flare stack and remained there through the last survey
burrows were observed then. (Table 5). A second one was observed at the stack in
mid-September. Surveys were sometimes limited by
reduced visibility, which may have resulted in reduced
Site fidelity tag detection.
Two days after tagging, divers sighted six tagged fish A total of 45 red snapper were released in the three
and found two detached tags on the bottom near the displacement experiments (see Table 6 for details), of
double webbing reef, indicating that tag loss has been a which 19 were tagged with T-bar tags and two with
problem. Of the remaining maximum of 19 tagged fish, sonic tags. During hauling of the cage, the fish were
fish re-sighted at the reef remained steady up to high ten noticeably distressed. They became bloated as a result of
in mid-August, after which the number dropped to one swimbladder expansion and turned in a downwardS48 I. Workman et al.
Table 6. Displacement experiments by date: number of fish
released (N), number tagged with T-bar (T) and sonic (S) tags, Experiment No. 1 (8 Aug. 1998)
and number of tagged fish relocated (S) or resighted (T) at the Rubble
capture location (R). Reef Displacement track
Return track
Date N T S R
N
08 Aug 1998 12 5 0 1 Displacement
09 Aug 1998 18 7 1 3 Horizontal: 0.37 km
10 Aug 1998 15 7 1 1
Vertical: 20 m
60
Experiment No. 2 (9 Aug. 1998)
50 Displacement
Number of fish
40 Horizontal: 0.43 km
Vertical: 21 m
30
N
Rubble
20 Reef
10
0
Mar-96 Apr-96 May-96 Jun-96 Jul-96 Aug-96
Figure 2. Red snapper counts by date at the double webbing Experiment No. 3 (10 Aug. 1998)
reef.
Displacement
Horizontal: 0.43 km (straight line)
swimming direction as the cage neared the surface. Vertical: 20 m
Upon release, they swam rapidly to the bottom. Descent N
Rubble
times ranged from 50 s to 1 min 14 s. Reef
After reaching the bottom, they remained more or
less stationary for 2–5 min before starting to swim.
Behaviour varied with each release. Fish in the first
release swam very close to the bottom and stopped three
times in an apparent effort to orient themselves before Figure 3. Displacement and return tracks for three horizontal-
the divers lost them at 7 min 50 s into the tracking part vertical displacement experiments (not to scale).
of the experiment. The sonic-tagged snapper in the
second experiment was separated from the other
fish from the first experiment and two from the second
released fish during descent. After reaching the bottom
were sighted on the rubble reef the day following their
and a 2 min 7 s stop, it spent most of its time swimming
releases.
between 1 m and 1.5 m off bottom, but occasionally
descended to the bottom where it would stop for a short
time and turn its left side toward the direction of travel. Discussion
Just before the divers had to come up, the fish swam
over another small rubble reef without hesitation or The comparative reef type study was of limited duration
change in heading. Fish of the third release, after but covered the primary recruitment period for age-0 red
starting to swim, made a big circle back to the point they snapper in the GOM (Moran, 1988). Also, the timing
started from, remained there for a short time, and then allowed observations on the interactions and behaviour
started swimming again. of age-0 and age-1 fish through the recruitment and
The course headings on all three releases corre- habitat transition phase and to interpret their habitat
sponded largely with the direction of the capture site preferences statistically.
(Figure 3). It took the sonic tagged fish about 25 min to Red snapper are categorized as reef fish and their
reach the capture site. On the two follow-up dives, divers reef dependency begins shortly after they leave their
recaptured the fish from the second experiment and planktonic life-stage (Szedlmayer and Howe, 1997;
picked up the signal of the snapper from the third Szedlmayer and Conti, 1999; Bailey, 1995). Habitat
experiment at the point of capture. One T-bar tagged requirements of the smallest settlers are apparently metHabitat preferences and site fidelity of juvenile red snapper S49
by the presence of small structures, including shells and materials that meet the needs of age-0 and age-1 red
burrows, but as they grow they prefer larger and more snapper might help to ensure increased recruitment to
complex structures (Bailey, 1995). However, recruitment adult habitats.
to these structures may be limited by the presence of
larger fishes. Our observations suggest that recruitment
of age-0 to the polyethylene webbing reefs is limited by Acknowledgements
the presence of age-1 snapper, because recruitment to
We thank Kendall Falana for his help in surveying the
the reef without the float and inhabited by less age-1 fish
reefs, John Mitchell for preparing figures and Deborah
was higher than to the reef with a float, which was
Seidel for her help in organizing the manuscript. We also
clearly preferred by age-1. Also, as age-1 started to move
express our appreciation to Drs Scott Nichols, Joanne
off the reefs, age-0 fish moved in. Finally, age-0 were
Shultz, and John Watson for their reviews and construc-
never observed at the double webbing reef, which was
tive criticism of the manuscript. We are indebted to the
occupied by age-1 throughout.
Mississippi Gulf Fishing Banks for deploying the con-
The proximity of large reef structures (natural as well
crete rubble reef for the attraction of juvenile red
as artificial reefs) to smaller structures may also influ-
snapper.
ence recruitment patterns. The first recruits appeared at
the 50-m reef set, almost a month prior to their appear-
ance at the 90-m reef set. Pre-settlement, planktonic References
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