Impacts of v-notching the American lobster.
KEY WORDS: v-notch, lobster, North Cape, oil spill, Homarus americanus
V-notching of ovigerous American lobsters (Homarus americanus H. Milne Edwards, 1837) has been practiced in the Gulf of Maine for decades (Daniel et al. 1989). A v-notch is a non-unique, identifying mark made by removing a small triangular piece of lobster exoskeleton and somatic tissue from one of 4 uropods (Fig. 1). A lobster with a v-notch is designated illegal for commercial sale until it is determined that the notch was lost as a result of the regrowth of the exoskeleton and somatic tissue. The goal of notching female lobsters is to delay harvest of known reproductive females for additional molt cycles, thereby increasing her likelihood of contributing eggs to the population while protected from harvest.
Using v-notches to delay fishing mortality of reproductive female lobsters has been increasing in popularity with commercial fishermen and fishery managers as an effective method to increase reproductive potential of exploited fisheries, and as an effective conservation strategy (Daniel et al. 1989, Tully 2001), but the method was not practiced in Rhode Island, or south of Cape Cod, prior to 2000. In this area, a recent development in management strategies, advocated by commercial fishermen and accepted by the Atlantic States Marine Fisheries Commission (ASMFC), is to v-notch nongravid sexually mature females. Despite the growing practice, there is little information that exists on the effects of survivorship and disease susceptibility of lobsters that have been v-notched. The duration of v-notch retention is also an unknown that relates directly to estimates of egg production resulting from delayed fishing mortality, and is an issue of increasing importance to lobster fisheries managers.
We designed an observational experiment to document survivorship, shell disease, and mark retention of a laboratory population of v-notched American lobsters. The experiment is a result of the North Cape oil spill and a need to understand the effects of v-notching and duration of mark retention to assess adequately the impacts of the North Cape lobster restoration efforts.
The oil spill that occurred in 1996 resulted in an estimated 9 million lobsters killed (Cobb & Clancy 1998, Cobb et al. 1999, French 1999). Based on a series of criteria (French 1999), it was estimated that approximately 1.248 million reproductive females (many reproducing multiple times) would be required to produce the estimated 17.2 billion eggs necessary to restore the original lobster population. The restoration strategy decided on in 2000 was to take 1.248 million reproductive females destined to market and place them back in the ocean over a period of years, after having received a v-notch. Although it has been illegal to harvest v-notched lobsters in federal waters since 1987 (Mauseth et al. 2001), the ASMFC Amendment 3 to the Interstate Fishery Management Plan of Lobster required the Rhode Island Marine Fisheries Council to pass regulations prohibiting possession and harvest of v-notched lobsters in 1998; Massachusetts, Connecticut, and New York followed with their own regulations shortly thereafter. To be legally protected, the v-notch was required to be 6.35 mm (1/4 in) in depth, without any setal hairs (Gibson & Angell 2006). The hope was that the delayed fishing mortality of reproductive v-notched lobsters would produce the 17.2 billion eggs required to meet the restoration goals (French 1999).
In southern New England, lobster shell disease has been recorded since the early 1980s (Estrella 1991). The prevalence of epizootic shell disease in wild lobsters increased considerably during the mid 1990s (Castro & Angell 2000, Glenn & Pugh 2005, Castro et al. 2006), affecting more than 45% of some Rhode Island areas by 2002 (Castro et al. 2006). The cause of lobster shell disease is still unknown, but increasing literature suggests environmental factors play a role (Robohm et al. 2005, Smolowitz et al. 2005, Glenn & Pugh 2006). Increased susceptibility to bacteria resulting from mechanical damage of the exoskeleton has been observed in other crustaceans (e.g., Cook & Lofton 1973), and concern over increasing bacterial exposure in lobsters resulting from v-notching has been studied (Getchell 1987); however, little information exists regarding increased susceptibility of v-notched lobsters to the current epizootic outbreak.
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One component necessary to analyze critically the success of the North Cape restoration program is to obtain independent data on variables related to mortality rate and increased disease susceptibility of notched lobsters, as well as the notch loss resulting from exoskeleton overgrowth. The objectives of this article are to describe the following parameters: (1) describe the survival of notched versus unnotched lobsters, (2) describe differences in the incidence of shell disease of notched and unnotched lobsters, and (3) describe the rate of notch loss (or overgrowth) with respect to molting frequency.
MATERIALS AND METHODS
Lobster collection and tank setup began approximately 1 mo before the start of the experiment, allowing for an approximate 2 to 4-wk adjustment period for lobsters to acclimate to tank conditions before being subjected to the stress of the experiment (Getchell 1987). Female lobsters were collected by Rhode Island Department of Environmental Management (RIDEM) lobster biologists during Rhode Island Lobster Research and Management Project sea sampling trips (trap caught) and were brought to holding tanks on the same day of capture. Lobsters collected for the experiment reflected the criteria for lobsters eligible for notching in the field; therefore, only female lobsters that were not egg bearing and had no visible signs of shell disease were chosen. The only exception to this was the size of the lobsters used in the experiment; collected lobsters were slightly under legal size (minimum carapace length (CL), 78.1 mm; maximum CL, 85.6 mm). All lobsters were housed at the RIDEM Marine Fisheries facility wet laboratory, Jamestown, RI, in rectangular tanks with flow-through ambient seawater coming from the lower east passage of Narragansett Bay. Ambient seawater was pumped directly into reservoir tanks located immediately above the lobster holding tanks. Reservoir and holding tank combined volume was approximately 700 gal. Flow was adjusted in each reservoir so water turnover rate was the same for all 3 tanks (1.3 times/hr). Temperature, salinity, and dissolved oxygen were monitored weekly.
Lobsters were housed in 3 tanks (tanks A, B, and C). Each tank was partitioned by a mesh barrier to create a control and experimental side. Twenty-five experimental lobsters were housed on one side of the barrier to represent an experimental group (E) and 15 control lobsters were housed on the other side of the barrier to represent a control group (C; tanks A-C, A-E, B-C, B-E, C-C, C-E), except for tank A,-C which only had 14 lobsters to start the experiment. Although the number of control and experimental lobsters was not equal, the partition dividing the 2 groups of lobsters was placed to create equal densities of lobsters on each side (7 lobsters/[m.sup.2]). Tanks were outfitted with PVC tubing and concrete blocks to provide sufficient dens and hiding places for all lobsters. To avoid injury to each other, rubber bands were used to close both chelae. Rubber bands were marked to identify individuals. Feeding was done on a daily basis during the warm-water months, and tapered off to 2-3 times per week during the cold-water months. Feeding was mainly squid (Illex illecebrosus) and butterfish (Peprilus triacanthus); however, Atlantic mackerel (Scomber scombrus), pogies (Brevoortia tyrannus), or blue back herring (Alosa aestivalis) were often used to vary diet. One week before the start of the experiment, a standard v-notch tool (used by North Cape v-notchers in the field) was used to mark each lobster in the experimental group with a v-notch in the right uropod next to the telson. V-notches were "double-tapped" in the same manner they would be by North Cape observers in the field (notched once, then notched in the same place again a few moments later, to remove any tissue extruding from the wound into the open space).
Observations began October 15, 2003, and occurred 1 day/ wk. Health of lobsters was recorded by noting the loss of any walking legs and noting whether each lobster was active or lethargic. Presence and degree of shell disease was monitored for each individual. The severity of shell disease was assessed by determining the degree of shell disease on the dorsal shell: none; low, <10%; moderate, >10 < 50%; severe, >50%. For experimental groups, v-notches were examined for the presence of setal hairs. When v-notch lobsters molted, v-notches were measured and classified as greater than or less than 6.35 mm (1/4 in) in depth (one of the management restrictions for a legal v-notch during the time of the North Cape restoration efforts). Lobsters were isolated in floating milk crates immediately prior to molting. All lobsters with soft shells were isolated until their shells were hard enough to return to the general population of their tank. On the final day of the experiment, September 16, 2005, all lobsters were recorded as usual and their tail flippers were photographed with a high-resolution digital camera (Canon Powershot; Canon USA, Inc., Lake Success, NY). All remaining lobsters were released in Fort Wetherhill Harbor, Jamestown, RI.
A repeated-measure analysis was performed to examine overall survival of control and experimental lobsters throughout the study. Survival fractions were calculated using the product limit or Kaplan-Meier method (Kaplan & Meier 1958). The test reports the variability of the fractional survival as an SE or 95% confidence intervals. SEs are calculated using the method of Greenwood (1926). Log-rank tests were used to compare the survival curves. When comparing 2 groups, the log-rank test is equivalent to the Mantel-Henszel test. This test generates a P value testing the null hypothesis that the survival curves are identical in the overall populations. Statistical significance was determined using an [alpha] = 0.05. The null hypothesis ([H.sub.0]) is that the treatments did not change survival.
Survival plots were used to obtain a general idea of the survival of control and treatment groups, and to be able to observe any significant changes in survival throughout the study period. Because of the small sample sizes, differences in precision for estimates in the tails, as opposed to the center of the distribution, were focused on more heavily (Lawless 1982).
V-notch retention was analyzed on a per-molt basis. V-notching of female lobsters in the field during the North Cape restoration efforts does not account for molt and reproductive condition (other than the lack of ovigerous eggs) of individual lobsters, and therefore is not a consideration in this analysis. Analysis assumes variation in intermolt stage at the time of the initial notch, and notch regrowth is analyzed as a function of the number of molt events until the v-notch crosses management definition thresholds, regardless of the timing of the molt event. V-notch retention was examined 2 ways. Notch size and presence of setal hairs were used independently and combined to categorize postmolt notches as being protected from harvest or not. Second, digital images of tail flippers photographed the final day of the experiment were analyzed, and remaining v-notches were measured to 0.1 mm. Photographed lobsters were categorized into groups depending on the number of times they molted since notched. A box-and-whisker plot was used to analyze remaining notch depth with respect to the number of molts, and to compare the remaining v-notch with past and present management definitions. V-notches were measured using image analysis software (Image Pro 4; Media Cybernetics, Silver Spring, MD). To standardize measurement of remaining v-notches, tangent lines ([L.sub.1]) were drawn connecting the highest points of the uropod on both sides of the open end of the V. A second line was run from the deepest point of the V (A), perpendicular to the tangent line. The notch was measured from point A to the 90[degrees] intersection of [L.sub.1] (Fig. 2).
From October 15, 2003, to September 16, 2005, 90 observations of control and experimental lobsters were performed to examine the effects of v-notching on the lobsters. Tank environmental condition (temperature, salinity, and dissolved oxygen) are summarized in Table 1. Environmental conditions were reflective of the parameters of the bay, and therefore fluctuated with season.
Survivorship and Shell Disease
Survivorship curves plotted for each tank compare the survivorship of control and treatment lobsters throughout the length of the study period (Fig. 3). No statistical differences in survival were observed for any of the study tanks (P values > 0.05). Median survival time for lobsters is undefined, because none of the lobster groups for any of the tanks ever reached a survival proportion less than 50%. Tanks A and B demonstrate almost identical survivorship between control and treatment lobsters throughout the entire duration of the study. Of particular interest is the hazard ratio at the start of experiment, immediately after the treatment lobsters are notched. No notched lobsters died in tank A immediately after notching, and only 1 lobster died in tank B immediately after notching.
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Although no statistical differences were found between control and treatment lobsters in tank C, it is important to note the difference in survivorship of those lobsters. Three (12%) treatment lobsters in tank C died immediately after being notched, before formal data recording began. Four additional lobsters in tank C died intermittently from the start of the experiment to the first summer (around week 35, Fig. 3C), when mortality of all lobsters in the experiment increased as a result of mortality associated with increased temperature (Table 1) and molting.
Shell disease of control and experimental lobsters was examined for differences in frequency and severity that v-notching may have caused. Figure 4 tracks the frequency and severity of shell disease throughout the length of the study. Although the severity of shell disease between control and treatment lobsters demonstrates some variability, the frequency or percentage of control and experimental lobsters with any degree of shell disease mimics each other with only minor variation (Fig. 4). Frequency of shell disease varied with season for all lobsters. Experimental lobsters typically demonstrate a lower percentage of shell disease overall than control lobsters, indicating that v-notching did not increase the frequency of shell disease in v-notched lobsters.
Sixty-three of the 75 experimental lobsters given v-notches survived to the first molt, and had tails that could be analyzed. Tails that could not be analyzed were those in which the notched uropod was completely eliminated or disfigured during a molt beyond what could reasonably be compared with a control lobster (5 could not be analyzed after first molt). After the first molt, 59% of v-notches were less than 6.35 mm (1/4 in) in depth (no longer protected as a result of notch size). By the second molt, 89% of notched lobsters had v-notches less than 6.35 mm (1/4 in) in depth (Table 2).
With regard to setal hair presence, 84% of notched lobsters had setal hairs after the first molt (no longer protected). After the second molt, 100% of the total lobsters notched exhibited setal hairs. When considering both variables that influence harvestability of lobsters, 97% of notched lobsters would have no longer been protected from harvest after the first molt because of either depth of notch or setal hair presence. After the second molt, 100% of notched lobsters were classified as harvestable (Table 2). The time from being notched to being harvestable ranged from 19M57 days (mean [+ or -] SE, 258 [+ or -] 13 days). Approximately 70% of lobsters became harvestable between 240 and 360 days (June to October, 2004) after being notched (Fig. 5).
Figure 6 is a box plot showing the remaining depth of v-notches in relation to the number of times the lobsters molted from the start of the study to the completion of the study. Of the lobsters that molted once (n = 5) during the study, the mean and median number of lobsters had v-notches less than 6.35 mm (1/4 in) in depth. All of those lobsters still had v-notches greater than 3.18 mm (1/8 in). Of the lobsters that molted twice during the study (n = 24), 100% of the lobsters had v-notches less than 6.35 mm (1/4 in) in depth, whereas only the 25th percentile had v-notches less than 3.18 mm (1/8 in) in depth. Only 1 lobster molted 3 times and was available for measurement at the end of the study; the notch measured 0.85 mm.
Survivorship and Health
There were no significant differences in survival of v-notched lobsters compared with those that were not v-notched. Despite the statistical evidence, it is important to use caution when examining these results because of the small sample size. Although control and experimental lobsters in tank C did not demonstrate significant differences in mortality, particular interest is paid to the period at the start of the experiment, when 12% of v-notched lobsters died immediately after notching. Examination of the survival trend lines indicate that, other than the mortality of v-notched lobsters in tank C at the start of the experiment, mortality rates were almost identical for each group throughout the entire length of the study. We believe it was not a tank effect causing the initial increased mortality in tank C, because the hazard ratio of individuals in tank C only diverged from the other two study tanks at the beginning of the study, then mimicked the other tanks exactly for the remainder of the study.
[FIGURE 4 OMITTED]
Trend lines of the percent of control and experimental lobsters with shell disease mimic each other almost exactly throughout the 2-y study period. It has been documented that environmental stress can increase disease susceptibility in lobsters (Robohm et al. 2005), and therefore was assumed here that tank conditions and affects (e.g., diet, crowding, stress) likely played a role in infection rate of tank-held lobsters (Malloy 1978, Getchell 1989, Prince et al. 1995). Although the rate and frequency of infection may be inflated compared with what would be observed in wild lobsters, no difference was observed between notched and unnotched lobsters, suggesting that the exposure (i.e., damage) created by notching did not increase rate or frequency of infection compared with control lobsters. Getchell (1987) conducted an experiment testing the vulnerability of v-notched lobsters to red-tail (Aerococcus viridans (vat.) homari) infection. The study performed histological observations of wound repair of v-notched uropods. In that study, it was witnessed that the wound was sealed by a heavy infiltration of hemocytes within 24 h. Observations made by RIDEM scientists performing separate wound retention-time experiments than ours in Jamestown, RI, came to similar conclusions (D. Costa, RIDEM, pers. comm., 2006). Despite the cause of shell disease, our results, in conjunction with these studies, suggests that v-notching does not impact the susceptibility, frequency, or severity of shell disease.
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To predict the reproductive potential of notching lobsters, it is imperative to understand how long the individual is protected from harvest and, therefore, be able to interpret how many times she will be able to produce eggs while protected. To do that, we attempted to quantify the retention time of v-notches in relation to the Rhode Island and southern Massachusetts legal definition of a v-notch while the North Cape project was in progress. At the time of the project, a lobster was protected from harvest as long as it had a v-notch greater than 6.35 mm (1/4 in) in depth and no setal hairs were present in the notch. We also compared v-notch fill-in rate with the new RIDEM v-notch definition 3.18 mm (1/8 in).
Our study suggests that when considering both v-notch size and setal hair presence, nearly 100% of lobsters are eligible for harvest after just 1 molt. The results show that this is clearly the result of the inclusion of setal hairs as a qualifying factor for harvest. When examining the two factors separately, after the first molt, 84% of notched lobsters exhibited setal hairs, whereas only 59% of notched lobsters had a v-notch less than 6.35 mm (1/4 in) in depth. After the second molt, no lobsters remained protected from harvest as a result of the lack of setal hairs, whereas 11% were still protected because their notch was greater than 6.35 mm (1/4 in) in depth. Results from this study indicated a relatively consistent duration of time from notched to harvestable. Although these results suggest that the majority of the experimental population was in similar stages of the molt cycle when notched, these results do not necessary translate to natural lobster populations. The results from this study more appropriately describe protection duration from a v-notch on a per-molt basis, as described later.
In scaling the restoration effort after the North Cape oil spill, Gibson (1998) estimated that a v-notch would provide 2 y of protection. Although this study was designed to account for the variability in life stage of notched lobsters, mimicking those lobsters notched in the field, an unknown in this experiment is the relative metabolic activity of our captive lobsters to similar lobsters in the natural environment. It has been proposed that food source and temperature are major factors that determine molt frequency and growth increment in lobsters (Chittleborough 1975, Mente & Houlihan 2002), and therefore will likely also affect v-notch fill-in rate. Many of our lobsters molted twice in 2 y, a rate faster than considered normal for female lobsters of this size (Herrick 1911, Aiken & Waddy 1976). The timescale, therefore, of the results may be compressed compared with lobsters in the natural environment. The results presented in this study assume that despite timescale compression of molt periodicity, the tissue regeneration on a per-molt basis is similar to that in the natural environment. The relative observations of the control and experimental lobsters would be expected to stay relevant; however, this result supports the need to examine v-notch retention in a population on a molt probability basis, rather than an intermolt duration.
Recently, southern New England fisheries managers have debated the legal definition of a v-notch. Our results indicate that because of the fill-in rate of v-notches, approximately 40% of lobsters will be protected for 1 molt with a v-notch definition of 6.35 mm (1/4 in), assuming setal hairs are disregarded. At this definition, our results indicate that almost no lobsters will still be protected after 2 molts. However, if the legal definition is reduced in half to 3.18 mm (1/8 in), regardless of setal hairs, then approximately 75% of the lobsters will still be protected from harvest after 2 molts.
This study suggests that v-notching the American lobster in Rhode Island waters does not have a significant effect on the lobster's survivorship or the frequency of shell disease. As a result of the inclusion of setal hairs in the v-notch definition at the time of the North Cape project, the majority of lobsters were only protected from harvest for 1 molt, unless the lobster was recaptured and renotched. If a v-notch definition of 3.18 mm (1/8 in), with or without setal hairs was in place, the majority of notched females would be protected for an additional molt, increasing her likelihood of contributing a second batch of eggs to the population before becoming eligible for harvest.
The authors thank the many people who assisted in tank maintenance and data collection throughout this 2-y experiment, especially John Holly, Elizabeth Kordowski, and Stephanie Hryzan. Thank you to Dr. Brian Quilliam for his assistance with statistical analysis and Dr. Steve Cadrin for his input into this manuscript. Thank you to the Rhode Island Department of Environmental Management for providing their services, wet lab, and tank space.
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BRYAN M. DEANGELIS, (1) * RICHARD COOPER, (2,4) MICHAEL CLANCY, (3) CHRISTOPHER COOPER, (4) THOMAS ANGELL, (5) SCOTT OLSZEWSKI, (5) WARREN (TED) COLBURN (4) AND JOHN CATENA (6)
(1) National Oceanic and Atmospheric Administration, Restoration Center, National Marine Fisheries Service, Narragansett Laboratory, 28 Tarzwell Drive, Narragansett, RI 02882; (2) University of Connecticut, 1084 Shennecossett Rd., Groton, CT 06340; (3) Eastern Connecticut State University, 83 Windham Street, Willimantic, CT 06226; (4) Ocean Technology Foundation, 34 Water Street, Mystic, CT 06355; (5) Rhode Island Department of Environmental Management, 3 Fort Wetherill Rd., Jamestown, RI 02835; (6) National Oceanic and Atmospheric Administration, Restoration Center, 55 Great Republic Drive, Gloucester, MA 01930
* Corresponding author. E-mail: Bryan.DeAngelis@noaa.gov
TABLE 1. Monthly mean, minimum (Min), and maximum (Max) temperature, salinity, and dissolved oxygen levels throughout the experiment. Temperature Salinity Dissolved Oxygen ([degrees]C) ([per thousand]) (mg/L) Date Mean Min Max Mean Min Max Mean Min Max Oct-03 15.7 14.6 16.8 30.1 29.2 30.9 6.8 6.4 7.2 Nov-03 12.6 11.3 14.6 30.9 30.8 30.9 7.8 7.2 8.5 Dec-03 7.7 6.3 9.5 30.3 29.7 31.0 9.5 8.8 9.9 Jan-04 5.2 3.9 6.4 28.9 27.4 30.6 10.0 9.7 10.5 Feb-04 2.7 2.6 2.9 29.6 28.8 30.7 11.4 11.0 11.7 Mar-04 3.7 3.2 4.4 31.2 30.0 32.2 10.6 10.0 11.1 Apr-04 6.5 4.6 8.3 30.8 30.0 31.6 15.9 8.4 91.6 May-04 12.3 10.2 14.3 30.1 29.2 31.2 7.8 6.9 8.1 Jun-04 15.0 14.4 15.8 31.0 30.8 31.2 6.9 6.5 7.2 Jul-04 18.7 17.0 19.9 31.2 30.5 32.0 6.5 6.5 6.5 Aug-04 20.0 19.5 20.3 31.7 31.5 31.8 6.4 6.3 6.6 Sep-04 20.0 18.6 21.3 31.3 31.0 31.4 5.9 5.5 6.2 Oct-04 15.7 14.0 17.6 31.0 30.5 31.6 7.0 6.4 7.4 Nov-04 12.0 10.5 13.5 31.0 30.5 31.6 7.8 7.4 8.2 Dec-04 8.3 5.9 9.8 30.0 29.4 30.2 8.9 8.4 9.7 Jan-05 4.7 2.6 6.2 30.3 29.5 31.3 10.9 9.0 12.9 Feb-05 2.6 2.0 3.0 29.1 28.8 29.4 12.2 11.4 13.8 Mar-05 4.1 3.4 4.6 28.8 28.7 28.8 11.2 11.2 11.3 Apr-05 7.7 4.8 9.5 28.8 28.7 29.0 9.9 8.5 11.7 May-05 10.4 9.6 11.4 30.3 29.2 31.4 7.9 7.6 8.1 Jun-05 15.2 13.6 16.6 30.0 29.3 30.4 6.8 6.3 7.5 Jul-05 18.6 17.4 19.9 30.5 30.3 30.8 6.1 5.6 6.6 Aug-05 20.3 20.2 20.5 30.7 30.6 30.9 5.7 5.3 6.0 Sep-05 20.3 20.0 20.6 31.1 31.0 31.1 5.6 5.4 5.9 TABLE 2. Rate of protected v-notch loss with respect to 2 notch criteria and successive molts. 1st Molt 2nd Molt Criteria for a Protected V-Notch % n % n Legal depth ([greater than or equal to] 6.35 mm) 41 26 11 5 Nonlegal depth (<6.35 mm) 59 37 89 39 No setal hairs 16 10 0 0 Setal hairs present 84 53 100 44 Not harvestable (both criteria) 3 2 0 0 Harvestable (both criteria) 97 61 100 44
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|Author:||Deangelis, Bryan M.; Cooper, Richard; Clancy, Michael; Cooper, Christopher; Angell, Thomas; Olszewsk|
|Publication:||Journal of Shellfish Research|
|Date:||Aug 1, 2010|
|Previous Article:||Physiological responses of brown crab (Cancer pagurus Linnaeus 1758) to dry storage under conditions simulating vitality stressors.|
|Next Article:||Mating and sperm storage of the Asian shore crab Hemigrapsus sanguineus.|