Observations of Juvenile Lobsters, Homarus americanus, on a Rock-Reef in Long Island Sound
Renee Mercaldo-Allen, Ronald Goldberg, Paul E. Clark, and Catherine A. Kuropat
Northeastern Naturalist, Volume 18, Issue 1 (2011): 45–60
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2011 NORTHEASTERN NATURALIST 18(1):45–60
Observations of Juvenile Lobsters, Homarus americanus,
on a Rock-Reef in Long Island Sound
Renee Mercaldo-Allen1,*, Ronald Goldberg1, Paul E. Clark1,
and Catherine A. Kuropat1
Abstract - Movements of juvenile Homarus americanus (American Lobster; hereafter
lobster) on and around a naturally occurring rock reef were monitored over a 3-year
period. Lobsters were sampled with baited traps deployed at each of ten sites. Catch-perunit-
effort (CPUE) and number of lobsters collected per trap haul was calculated for each
sampling event. Physical habitat, visually characterized by underwater video and diver
observations, differed among sites. Lobster CPUE was significantly greater at rocky sites
(>70% density of cobble and/or boulder) containing complex structure, vertical relief
from the seafloor, and colonies of macroalgae, sponge, and hydroids. Lobster CPUE was
highest from late June to mid-July. Lobsters ranged from 18 to 82 mm carapace length
(CL), with 90.7% of tagged lobsters measuring between 30 to 60 mm CL. Relative lobster
abundance remained similar over the course of the study. Catch data were kriged to
illustrate spatial patterns of distribution. Over the study period, a total of 934 lobsters
were tagged and 66 were recaptured, for an overall recapture rate of 7.1%. The majority
of recaptured animals (88%) were found at the original tagging site or adjacent sites, with
one lobster remaining at liberty for 397 days. Most juvenile lobsters showed fidelity to
their initial site of capture on a small, relatively isolated patch of rock-reef habitat in the
central basin of Long Island Sound.
Introduction
Homarus americanus H. Milne-Edwards (American Lobster; hereafter lobster)
is a commercially and recreationally important decapod crustacean with a
range from Labrador to North Carolina (Holthuis 1991). Long Island Sound is
at the southern end of the inshore shallow-water distribution for this species and
has historically supported a commercial fishery. During fall of 1999, a largescale
mortality of Long Island Sound lobsters resulted in a significant population
decline (CT DEP 2000). Annual harvests from the commercial lobster fishery
currently remain well below pre-mortality levels (Giannini and Howell 2007).
Research efforts, initiated in response to the die-off, suggest that lobsters were
weakened by adverse environmental conditions (e.g., high seawater temperatures,
low dissolved oxygen levels, elevated sulfide, and/or ammonia), and may
have become susceptible to infectious pathogens (Pearce and Balcom 2005).
This widespread mortality and subsequent economic loss in Long Island Sound
has highlighted the need for a better understanding of the habitat requirements of
lobsters in this region and their potential for stock recovery.
1NOAA Fisheries Service, Northeast Fisheries Science Center, Milford Laboratory, 212
Rogers Avenue, Milford, CT 06460. *Corresponding author - renee.mercaldo-allen@
noaa.gov.
46 Northeastern Naturalist Vol. 18, No. 1
The role of shelter-rich habitat in the benthic ecology of the American Lobster
has been well described (see reviews by Barshaw and Lavalli 1988, Cooper and
Uzmann 1980, Lawton and Lavalli 1995, Mercaldo-Allen and Kuropat 1994).
Rock-reef habitats, comprised of cobble and boulders, provide multi-dimensional,
shelter-rich environments for lobsters in coastal southern New England (Steimle
and Zetlin 2000) at all ontogenetic stages (Cobb et al. 1999). This habitat may be
critical for lobster survival during the early years of life, when recruitment and
abundance are closely tied to shelter availability (Cobb et al. 1999, Langton et al.
1996, Wahle 1992, Wahle and Incze 1997, Wahle and Steneck 1991). Composition
of substrate along the seafloor determines structural complexity and shelter
availability within habitats and can influence the density, biomass, and size
structure of lobster populations (Beck 1995, Hudon 1987, Incze and Wahle 1991,
Wahle 1993, Wahle and Steneck 1991).
Studies have documented distribution, abundance, and movement of lobsters
within Long Island Sound (DRS 2005, 2006; Howell et al. 2005; Lund et al.
1973), but specific fine-scale habitat use by juvenile lobsters in this region has
not been well studied. Within the central basin of Long Island Sound, rock-reef
habitat occurs in isolated patches and constitutes a relatively small portion of the
overall available seafloor (Poppe et al. 2000). Discrete patches of shelter-rich
habitat scattered amid featureless bottom have been shown to harbor resident
populations of juvenile lobsters (Briggs and Zawacki 1974).
The data presented here were collected coincidentally during a project investigating
finfish settlement on a small reef in the central basin of Long Island Sound.
The presence of significant numbers of juvenile lobsters in this habitat led us to
initiate a tagging study to determine site fidelity and relative lobster abundance
within this small, relatively isolated patch of rock-reef habitat.
Methods
Study area
Our study area was a natural rock and cobble reef, located in Long Island Sound
off the coast of Milford, CT near Charles Island at approximately 41°11'13.73"N
and 73°3'48.65"W (Fig. 1). The reef covers an area of 0.25 km2, is discontinuous,
and varies in rock size and density. Seawater depth varies from 3 to 4 m at low
tide, with a 2-m tidal range, and is often highly turbid from resuspension of sediments
by waves and tidal currents.
Rock size was classified nominally with the Udden-Wentworth grain-size
scale (Lewis and McConchie 1994) as pebbles (4–65 mm), cobble (>65–250
mm), and boulders (>250 mm–1 m). Physical characteristics of each site were
assessed from Smith-McIntyre benthic grab samples, SCUBA diver observation,
and from video and still images. Sediment samples from the benthic grab were
sieved and characterized grossly. The Sea Boss System (Blackwood et al. 2000),
developed and operated by the United States Geological Survey, Woods Hole Science
Center, provided video and still seafloor images. Descriptions of sites were
based on evaluations of images from still photos and video footage.
2011 R. Mercaldo-Allen, R. Goldberg, P.E. Clark, and C.A. Kuropat 47
Figure 1. The rock-reef study and study sites located off Charles Island on the coast of
Milford, CT in the central basin of Long Island Sound. Benthic substrate on the study
sites was characterized as: 70% boulder at sites 4, 5, and 10; < 50% boulder at site 3;
30% cobble at site 2 and 10% cobble at site 7; featureless silt-sand at sites 1, 6, and 8;
and mud at site 9.
48 Northeastern Naturalist Vol. 18, No. 1
Bottom sediments in the study area ranged from all silt-clay to a mixture of
silt-clay and sand. Sediment was covered with discontinuous patches of shell
hash or pebbles and a fine silt layer. Reef sites 4, 5, and 10 were composed of
over 70% dense boulder cover, providing vertical relief up to 1 m off the seafloor.
Cobble and boulder at these sites were heavily encrusted with epibiotic growth
including seaweed, sponges, hydroids, and bryozoans. Density of boulder cover
at site 3 was less than 50%. At sites 2 and 7, only 30% and 10% of the bottom,
respectively, was covered by cobble. Sites 1, 6, and 8 were characterized by a
featureless silt-sand bottom covered with high densities of Crepidula fornicata
L., shell hash, and low vertical relief. Site 9 (2004 and 2005 only) was primarily
a mud bottom.
Ten sites were established, with five located on reef structure (sites 2, 4, 5,
7, and 10) and five sites situated in areas with little to no reef structure (sites 1,
3, 6, 8, and 9). Sites were marked with a cinder block and attached buoy, and
latitude and longitude were recorded using shipboard GPS. During 2006, site 9
was dropped from the study to accommodate commercial clamming activities.
Sampling took place from either the Milford Laboratory’s 15-m NOAA R/V Victor
Loosanoff or a smaller 7-m vessel.
Lobster sampling
Sampling was conducted using commercially available, wire-frame 3-mm
nylon mesh fish traps. The traps measured 23 × 23 × 46 cm and contained no
escape vents. Each double-entry single-chambered trap had a flexible 25-mmdiameter
ring entrance opening at each end and a 5-kg steel plate along the
base to assure stability. Three traps were set at each site in 2004. Since this
study was initially designed to sample young fish, the presence of lobsters in
the traps was unexpected. For the second and third study year, we increased
sampling effort from 3 to 6 traps per site. Prior to deployment, traps were
baited with clam meats.
Traps were deployed from June through October of 2004, June through September
2005, and May through November 2006. Sampling periods varied with
study year as a result of vessel availability and weather conditions. Traps were
checked daily, although infrequently, traps soaked for more than a week due to
weather. All sampling was conducted during daylight hours between 0800 and
1400 h. A total of 46, 41, and 51 sampling trips were completed in 2004, 2005,
and 2006, respectively.
Upon retrieval, lobsters were removed from a trap, carapace length (CL) was
measured to the nearest millimeter with a caliper, and sex was noted. Lobsters
>30 mm were marked with a stainless steel sphyrion anchor-style tag (Floy
Tag and MFG., Inc.) made of polyolefin tubing. Uniquely numbered tags were
inserted into the dorsal musculature between the cephalothorax and the first abdominal
segment, to one side of the midline, using the tip of a 5-cc (21 gauge ×
1.5") disposable syringe (Smith et al. 2001) to allow identification and tracking
of individual animals. Sphyrion tags have been widely used in mark-recapture
2011 R. Mercaldo-Allen, R. Goldberg, P.E. Clark, and C.A. Kuropat 49
studies with American Lobsters (e.g., Campbell 1989, Scarratt and Elson 1965).
However, tag-retention rates vary from 10 to 36%, as tags can be lost during
molting (Cooper 1970, Ennis 1986). Lobsters which were damaged or had recently
molted were released without tagging. Animals which measured <30 mm
in size were considered too small to tag and were also released. Sampling was
restricted to the reef study area, and recaptures outside this zone included only
incidental returns reported by commercial lobstermen.
Near-bottom seawater temperature (°C), salinity (PSU), and dissolved oxygen
(mg/L) were measured with a handheld YSI meter (model 88) at two sites (6 and
10) during each sampling trip. An automated water temperature logger (Onset
HOBO Water Temp Pro) was attached to the top of one trap deployed at site 5
during the 2005 and 2006 sampling seasons.
Statistical procedures
Catch-per-unit-effort (CPUE) was calculated as number of lobsters caught per
individual trap haul. The CPUE data could not be transformed to meet ANOVA
assumptions of normality and homogeneity of variance, so a non-parametric
Kruskal-Wallis ANOVA on ranks was used. Tukey’s multiple comparisons were
used for post-hoc testing to identify differences in relative lobster abundance
among sites within each year. Pearson correlation was used to evaluate the
relationship between lobster CPUE and both trap soak time and weekly mean
seawater temperature. Spatial extrapolations of lobster distribution and relative
abundance were generated by ordinary kriging for 2004 and 2005, using the
Geospatial Analyst extension of ARC GIS 9.2 software. Use of a spherical model
was validated by estimation of model strength, based on prediction-error plots
generated by the software.
Results
Abundance, distribution, and size composition
Size-frequency distributions for lobsters at each study site by year are
shown in Table 1. The smallest lobsters collected in our traps measured 18
mm CL—a size at which they transition from a shelter-dependent lifestyle
to a free-roaming existence (Hudon 1987, Lawton 1987, Lawton and Lavalli
1995). Due to their cryptic nature and strong association with the benthos,
lobsters below this size have not been successfully sampled by trapping (Sclafani
and Smith 2003). Our traps collected primarily juvenile lobsters (30 to 60
mm CL); however, occasionally larger Lobsters entered the traps due to the
flexibility of the traps.
Lobster CPUE for each site by year is shown in Figure 2. In all three years,
lobsters were significantly more abundant at reef sites 4 and 5 compared to
non-reef sites 1, 2, 7, 8, and 9 (H = 131.602 [2004], H = 214.882 [2005],
H = 196.416 [2006], df = 9, P < 0.001). There were no differences in lobster
abundance between reef sites 4, 5, and 10 in all years. Lobsters were significantly
more abundant at reef sites 4 and 5 than at non-reef site 6 during 2004
50 Northeastern Naturalist Vol. 18, No. 1
and 2006. Lobsters were more abundant at reef site 4 than at non-reef site 3 in
all three years. Significantly more lobsters were found at reef site 5 compared
to non-reef site 3 during 2005. Reef site 10 had significantly more lobsters than
non-reef site 9 during 2005. Neither trap soak time (r = 0.025, P = 0.0409) or
weekly mean seawater temperature (r = -0.033, P = 0.0154) correlated significantly
with lobster CPUE.
Lobster CPUE was highest from late June to early July in all years. Ambient
seawater temperatures and Lobster abundance for each week of sampling by year
are also shown in Figure 3. Seawater temperatures reached maximum values
between late July and early August. Near-bottom salinity ranged from 24 to 29
PSU, and dissolved oxygen levels never dropped below saturation.
Size-frequency distribution for all study years combined is shown in Figure 4.
The majority of lobsters measured between 30 and 60 mm CL, while the total size
distribution ranged from 17.9 to 82.4 mm CL. Sex ratios (female:male) measured
1.02 for 2004 and 2005 and 0.85 for 2006.
Ordinary kriging of Lobster CPUE data illustrates spatially extrapolated
lobster abundance in the vicinity of the reef for 2004 and 2005 (Fig. 5; the 2006
Table 1. Size (mm CL)-frequency distributions for American Lobsters collected from sites 1–10
during 2004, 2005, and 2006. ND indicates no data.
Site
Year CL 1 2 3 4 5 6 7 8 9 10
2004 11–20 0 0 0 0 0 0 1 0 0 0
21–30 0 2 3 13 4 3 3 4 3 1
31–40 2 3 6 31 22 3 3 11 2 10
41–50 2 2 8 32 15 6 1 2 3 6
51–60 0 4 1 32 14 2 0 1 4 6
61–70 1 1 1 9 3 0 0 0 3 0
71–80 1 1 0 6 0 0 0 0 1 0
81–90 0 0 1 1 0 0 0 0 0 0
ND 0 1 8 19 13 7 3 1 5 7
2005 11–20 0 0 0 0 1 0 0 0 0 0
21–30 3 3 1 6 8 9 3 3 1 6
31–40 5 7 7 28 40 19 3 9 3 23
41–50 7 9 13 40 41 11 3 4 2 16
51–60 1 7 10 37 16 5 3 2 2 11
61–70 1 2 2 10 1 1 0 0 2 3
71–80 0 0 1 0 1 0 0 0 0 0
ND 0 0 0 0 2 1 0 0 0 0
2006 11–20 0 0 0 0 0 0 1 0 ND 0
21–30 0 0 5 3 4 2 3 2 ND 4
31–40 2 5 8 33 30 13 5 5 ND 14
41–50 4 7 9 27 30 9 6 8 ND 16
51–60 0 3 5 24 18 1 5 2 ND 13
61–70 1 0 3 4 2 1 1 0 ND 1
71–80 0 0 0 0 0 0 0 0 ND 0
2011 R. Mercaldo-Allen, R. Goldberg, P.E. Clark, and C.A. Kuropat 51
Figure 2. Comparison of lobster CPUE (catch/trap haul) at the ten sampling sites is shown
for each year of the study (2004, 2005, and 2006). Mean CPUE are plotted with error bars
representing standard error. Common superscript letters above error bars denote statistically
similar groups (P > 0.05). ND indicates no data.
52 Northeastern Naturalist Vol. 18, No. 1
Figure 3. Lobster CPUE (catch/trap haul) versus bottom seawater temperatures over the
sampling season for each study year (2004, 2005, and 2006). Solid black circles connected
by lines represent temperature. ND indicates no data were collected during that month.
data was not included since site 9 was dropped). Lobsters were most abundant
on the eastern limb of the reef, corresponding generally to the darker bands on
the charts near sites 4, 5, and 10. These sites had the highest density of cobble
and boulder covered with epifauna and represented the greatest habitat complexity.
Fewer lobsters were sampled at all other sites that had lower densities
of cobble and boulders.
2011 R. Mercaldo-Allen, R. Goldberg, P.E. Clark, and C.A. Kuropat 53
Mark-recapture
Table 2 shows mark-recapture data for 2004, 2005, and 2006. A total of 934
lobsters were tagged. During 2004, 19 of 279 tagged lobsters were recaptured
after remaining at liberty from 4 to 105 days. Seventeen of these lobsters were
recaptured at the original tagging site, and two Lobsters were recaptured twice.
During 2005, 28 of 368 lobsters were recaptured after 5 to 67 days at liberty.
Twenty-four were recaptured at the tagging site, and one lobster was recaptured
twice. Nineteen of 287 were recaptured in 2006 with 17 recurring at the original
tagging location, and days at liberty ranged from 1 to 397. Over the entire study,
most of the recaptured lobsters (88%) were found at or adjacent to their original
tagging sites. Among eight lobsters tagged during 2004 and 2005 and recaptured
1 year later, six were collected at their tagging sites.
Discussion
Traps have limitations as quantitative sampling devices for lobsters. The
probability that a lobster will enter a trap varies with molt stage, reproductive
cycle, trap design, gear saturation, choice of bait, gender, behavioral
interactions, size, lunar and diurnal cycles, temperature, and hydrographic
conditions (Jury et al. 2001, Miller 1990). Underwater observations show that
Figure 4. Size-frequency distribution of lobsters collected at the ten sampling sites from
2004 through 2006. No data was available from site 9 during 2006.
54 Northeastern Naturalist Vol. 18, No. 1
Figure 5. Extrapolated geospatial patterns of distribution and abundance, based on ordinary
kriging of the total number of American Lobsters sampled for 2004 and 2005, at
each of the 10 sampling locations. Color gradations indicate the model estimate of the
relative number of lobsters in that band. Scale values denote the largest number of lobsters
in each band.
2011 R. Mercaldo-Allen, R. Goldberg, P.E. Clark, and C.A. Kuropat 55
many more lobsters approach a trap than actually enter and that trap catches
may underestimate true population density of lobsters in the trapping area (Jury
et al. 2001). Estimates of CPUE may be affected by undetected movement of
small lobsters in and out of traps. Despite these shortcomings, traps offer an
inexpensive, easily deployed method of continuous sampling that allows a
relative assessment of population dynamics in a discrete area (Dunnington et
al. 2005). Although a trap-based lobster CPUE does not represent an absolute
measure of lobster density (Jury et al. 2001), it does provide an index of relative
abundance (Scheirer et al. 2004).
We found higher relative abundance of lobsters on this relatively isolated
complex rock habitat as compared to adjacent areas with less structure. The
majority of lobsters were collected (66% of all captured and 71% of those recaptured)
from two sites with the highest concentrations of cobble and boulder
substrate. Significantly fewer lobsters were found on open silt-clay or sand bottom
locations having little cobble or boulder. Kriging of the CPUE data illustrates
consistency of this trend during 2004 and 2005, where the highest concentrations
of lobsters were associated with high rock density. The limited availability of
Table 2. Lobster size (mm CL)-frequency distributions for tagged and recaptured lobsters collected
from sites 1–10 during 2004, 2005, and 2006. ND indicates no data.
Year Lobster size (CL) Number tagged Number recaptured Recapture rate (%)
2004 21–30 21 0
31–40 89 0
41–50 75 5
51–60 62 10
61–70 17 2
71–80 9 2
81–90 2 0
ND 4 -
Total 279 19 6.8
2005 21–30 0 0
31–40 125 1
41–50 135 13
51–60 84 12
61–70 19 2
71–80 2 0
ND 3 -
Total 368 28 7.6
2006 21–30 1 0
31–40 102 4
41–50 103 4
51–60 65 9
61–70 13 2
71–80 0 0
ND 3 0
Total 287 19 6.4
Total 934 66 7.1
56 Northeastern Naturalist Vol. 18, No. 1
rock habitat in central Long Island Sound could influence population size and
recruitment potential for lobster. Cobble and boulder patches are relatively uncommon,
representing a very small percentage of the total area of the central
basin. Typical surficial sediments in Long Island Sound consist of sand along
the nearshore margins and silty-clay in the low-energy environment of the basins
(Poppe et al. 2000). The sand substrate beneath the boulder and cobble may have
provided additional refuge for lobsters, which can modify or create shelter by
excavating or bulldozing soft sediment (Barshaw and Bryant-Rich 1988, Cobb
1971, Miller et al. 2006, Wahle 1992).
Lobster abundance was highest during late June and early July when seawater
temperatures ranged from 18 to 22 °C. Studies in eastern Long Island Sound
also reported the highest mean CPUE of Lobsters at this time (DRS 2005). We
observed a decline in Lobster CPUE during mid-to-late July at peak seawater
temperatures (22–24 °C). Lobsters experience stress at bottom temperatures
exceeding 20 ºC (Howell et al. 2005, Pearce and Balcom 2005), and may be less
likely to forage for food or enter traps at elevated water temperatures. Since the
relationship between lobster catch and temperature can vary geographically and
over time, it can be difficult to link abundance with environmental conditions
(Koeller 1999).
Our overall recapture rate of 7.1% (66 recaptured / 934 tagged) was similar
to that found by Dunnington et al. (2005), who recaptured 8.6% (1023) of
11,856 lobsters tagged in Maine, and Watson et al. (1999), who recaptured
10.9% (1212) of 11,143 lobsters tagged in New Hampshire. During our study,
juvenile lobsters showed fidelity to the sites on the rock reef where they were
tagged with little evidence of movement among sampling sites. The majority
of recaptured lobsters (88%) were found on the reef up to a year after tagging.
When a tagged lobster was recaptured at another location, it was almost
always at an adjacent site, fewer than 10 m away. In Long Island Sound,
movement of lobsters appears to be localized and nonmigratory (Briggs and
Mushacke 1984; Briggs and Zawacki 1974; Cooper 1970; DRS 2005, 2006;
Howell et al. 2005; Lund et al. 1973). Most lobsters collected in our traps were
below legal harvest size, reproductively immature, and would not be expected
to undergo the deep-shallow water movements undertaken by larger mature
animals, which migrate in response to seasonal temperature cues (Campbell
1989, Campbell and Stasko 1986). Recapture of seven of eight lobsters on
the reef nearly a year or more after tagging, suggests that lobsters overwinter
on the reef or return in spring following fall/winter migration to deeper more
stable waters. More than half of small (50–59 mm CL) resident lobsters in a
shallow Massachusetts cove remained there during winter (Karnofsky et al.
1989) and similar overwintering behavior was observed in lobsters inhabiting
artificial pumice concrete shelters in otherwise open bottom near Point Judith
Rhode Island (Sheehy 1976). Also, small ovigerous lobsters (less than 93 mm CL) in
the Gulf of Maine were found to remain in shallow waters year-round (Cowan
et al. 2007).
2011 R. Mercaldo-Allen, R. Goldberg, P.E. Clark, and C.A. Kuropat 57
Our study suggests that this rock-reef habitat concentrates and supports local
populations of lobsters and may provide important ecological services to juvenile
lobsters. This may be of particular importance to the commercial fishery in Long
Island Sound, where lobster harvests have markedly declined and catches remain
well below historic levels. Identification of those habitats that support lobster
populations may assist in developing measures to enhance recovery of this species.
Our study suggests that even a relatively small and isolated patch of rock
reef may provide stable and valuable habitat for juvenile lobsters in the central
basin of Long Island Sound
Acknowledgments
The authors thank Captain Robert Alix and Werner Schreiner of the R/V Victor Loosanoff
for field support; Brian Hooper, Jose Pereira, Dylan Redman, James Reidy, Ann
Marie Salvato, George Sennefelder, Lauren Vinokur, and John Ziskowski for technical
assistance; Larry Poppe, Dan Blackwood, and Ivar Babb for video imagery; Larry Williams
for allowing us to place our traps on his leased shellfish grounds; and Barry Smith,
Mark Dixon, Dave Veilleux, and the rest of the NMFS/Milford Laboratory dive team for
underwater photography and habitat assessments. Use of trade names does not imply
endorsement by the NOAA Fisheries Service.
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