<I>Panulirus Marginatus,</I> and Slipper Lobster

BULLETIN OF MARINE SCIENCE. 89(2):529–549. 2013 http://dx.doi.org/10.5343/bms.2012.1047

Annual and Long-term Movement Patterns of Spiny Lobster, Panulirus marginatus, and Slipper Lobster, Scyllarides squammosus, in the Northwestern Hawaiian Islands

Joseph M O’Malley and William A Walsh

abstract Crustaceans exhibit diverse movement behaviors, which can complicate sustainable management if poorly understood. Annual and long-term movement patterns by Hawaiian spiny lobster, Panulirus marginatus (Quoy and Gaimard, 1825), and scaly slipper lobster, Scyllarides squammosus (H. Milne-Edwards, 1837), at four locations in the Northwestern Hawaiian Islands (NWHI) were estimated by using tag/recapture methods. Both species exhibited very limited movement, moving <1 km from the tagging site. Even after 5 yrs at liberty, the mean distance moved from the original tagging location was 0.09 and 0.23 km for male and female P. marginatus, respectively, and 0.54 and 0.35 km for male and female S. squammosus, respectively. Although 1% of P. marginatus and 2.8% of S. squammosus moved >5 km, there was no evidence of regular long-distance migrations or directed movements. Sex and location significantly affected distance moved by both species; however, there was no clear pattern. Size-at- tagging was a significant factor for S. squammosus, for which distances moved varied directly with increasing size; however, this is not necessarily indicative of large-scale unidirectional movements and may simply indicate that larger S. squammosus have larger home ranges. The lack of large-scale movements of both species probably reflects habitat characteristics. The NWHI do not provide the typical habitats juvenile lobsters require, there are no large seasonal temperature changes, and contranatant migrations are unnecessary because newly hatched larvae have access to offshore currents. The small home ranges of these species suggest that marine protected areas may be a viable conservation option.

Decapod crustaceans exhibit a wide range of movement behaviors. Herrnkind (1980) classified these movements, based on spatial extent and behavioral patterns as (1) homing, (2) dispersive, and (3) migratory. Palinurid and scyllarid lobsters exhibit all three types, sometimes more than one during an individual’s life span [e.g., Panulirus argus (see Appendix 1 for species authorities; Kanciruk and Herrnkind 1978, Davis and Dodrill 1989)] and the pattern can vary spatially within a species [e.g., Scyllarides latus (Lavalli et al. 2007)]. This diverse array of movement patterns can complicate management if a species’ behavior is unknown. Understanding lobster movements sheds light on recruitment processes, spatial variability in life history, and fisheries catch data. It also directly affects conservation decisions such as large-scale spatial management (Goethel et al. 2011), marine protected areas (Moffitt et al. 2011), and pres- ervation of nursery habitats (Butler et al. 2005). The Hawaiian Archipelago consists of the main Hawaiian Islands (Hawaii to Niihau) and the Northwestern Hawaiian Islands (NWHI). The latter are a series of islands, reefs, seamounts, and atolls (hereafter referred to as banks) extending

Figure 1. Map of the Hawaiian Archipelago, including the Northwestern Hawaiian Islands. Study sites are Necker Island, Gardner Pinnacles, Maro Reef, and Laysan Island. Contour of main Hawaiian Islands represents land. Contour of Northwestern Hawaiian Islands represents 20 fathom curve. Base bathymetric map was modiﬁed from Paciﬁc Islands Benthic Habitat Mapping Center, School of Ocean and Earth Science and Technology, University of Hawai‘i at Mānoa by A Andrews (NOAA/PIFSC). approximately 2000 km across the subtropical Pacific F( ig. 1). These banks are characterized by extensive shelves <40 m deep and defined by steep drop-offs to deeper water (Parrish and Polovina 1994). The shallow shelves provide essential habitat for the endemic Hawaiian spiny lobster, Panulirus marginatus, and the scaly slipper lobster, Scyllarides squammosus (Uchida et al. 1980). A commercial fishery targeted both these species from 1976 to 1999, and despite its becoming Hawaii’s most valuable demersal fishery D( iNardo and Marshall 2001), little is known about the behavior of either species, including movement patterns both among and within banks. The channels between NWHI banks exceed 1000 m (Uchida and Uchiyama 1986), which is beyond the typical depth range of Panulirus (Butler et al. 2006) and Scyllarides (Spanier and Lavalli 2006). This deep water has been assumed to prevent movement of either species between banks (Parrish and Polovina 1994). Genetic studies indicated a single-stock of P. marginatus throughout the archipelago (Shaklee and Samollow 1984, Seeb et al. 1990), presumably connected only by larval dispersal because of the inability of lobsters to traverse the deep water between banks. However, between-bank movements have never specifically been examined for either species. Historically, within-bank NWHI P. marginatus movements patterns have been deduced from observations and catch data. Queuing behavior of P. marginatus has been recorded at Kure Atoll, NWHI, and Kaneohe Bay, Oahu, but each of the four diver-observed events consisted of only two adult lobsters moving across o’malley and walsh: NWHI lobster movements 531 open sandy areas between reef patches (MacDonald et al. 1984). Panulirus marginatus was suspected to migrate across Necker Island based on size-frequency distributions derived from the National Oceanic and Atmospheric Administration (NOAA) Fisheries annual NWHI fisheries-independent lobster surveys. Relatively small lobsters were captured in the northern portion of the bank, so it was assumed this area contained habitat necessary for juveniles (DiNardo and Marshall 2001). A higher proportion of larger P. marginatus found outside this putative nursery area was thought to occur because of this species’ migrations to adult habitat at the southern end of the bank. Despite the lack of lobster movement and Necker Island habitat information, this assumption was sufficiently convincing to lead to a revision of the NOAA Fisheries research survey sampling design. To date, there is no knowledge about NWHI S. squammosus within-bank movements. The objectives of this study were to estimate annual and long-term movements and describe between- and within-bank movement patterns of NWHI P. marginatus and S. squammosus using tag and recapture methods. In addition, we examined the effects of several factors expected to influence movements of these species: sex, size, and location.

Materials and Methods

Locations and Dates.—Lobster tag/recapture operations were conducted aboard two commercial fishing vessels, each of which was chartered for 30 d per annum and cap- tained by an experienced NWHI lobster fisherman. Operations occurred at Necker Island (23°30Ń, 164°35´W; Fig. 2A), Gardner Pinnacles (25°00Ń, 168°50´W; Fig. 2B), and Maro Reef (25°30Ń, 170°45´W; Fig. 2C), the banks where the NWHI commercial fishery was historically concentrated, and Laysan Island (25°45Ń, 171°45´W; Fig. 2D), which had always been closed to fishing except in 1986. In 2003, annual tagging began with P. marginatus at Necker Island and S. squammosus at Maro Reef. During 2004 and 2005, both species were tagged at Necker Island and Maro Reef. In 2006, tagging expanded to Gardner Pinnacles and Laysan Island; during 2006–2008, tagging took place at all four banks. The tag/recapture cruises were conducted between June and September of each year. Within each bank, effort was concentrated in areas with greater lobster densities, as indicated by habitat (depth and sand vs hard bottom) and based on the captain’s previous lobster fishing experiences in the NWHI. Trapping, however, took place throughout the entirety of each bank to capture individuals that moved greater distances from their release locations (Fig. 2A–D). The sole exception to this was at Gardner Pinnacles where only the northern half of the bank was sampled. Depth of fishing ranged from 17 to 135 m, with the majority (89%) of traps set in depths <37 m. Capture, Tagging, and Recapture.—Lobsters were captured and recaptured with standard commercial molded black polyethylene traps (Fathoms Plus, San Diego, California), which are dome-shaped and single-chambered, with two entrance cones (dimensions = 980 × 770 × 295 mm, inside mesh dimensions = 45 × 45 mm). Mesh paneling was placed over the escape vents to prevent the escape of small lobsters, thereby maximiz- ing the size range of captured lobsters. Fifteen strings, consisting of 20 traps/string, were fished overnight. Traps were spaced 36.6 m apart on the strings and baited with 1 kg of Pacific chub mackerel, Scomber japonicus. Passive integrated transponder (PIT) tags [Destron Technologies, South St. Paul, Minnesota (tag model TX1411SST, dimensions: 12.50 × 2.07 mm); Trovan, Ltd., UK (tag model 100A, dimensions: 11.5 × 2.2 mm)] were injected into the distal tail segment (O’Malley 2008) and detected with a Destron Technologies Model 2001F-ISO portable 532 BULLETIN OF MARINE SCIENCE. VOL 89, NO 2. 2013

Figure 2. Lobster trapping effort by year at (A) Necker Island, (B) Gardner Pinnacles, (C) (Opposite page) Maro Reef, and (D) Laysan Island. Each point represents a string of 20 traps with year represented by different symbols. o’malley and walsh: NWHI lobster movements 533

Figure 2. Continued. 534 BULLETIN OF MARINE SCIENCE. VOL 89, NO 2. 2013 transceiver (Digital Angel Corporation, St. Paul, Minnesota). PIT tag loss was assumed negligible based on the results of a double-tag experiment that tested streamer tag and PIT tag retention rates in NWHI P. marginatus (O’Malley 2008). An important detail of this study is that because the commercial fishery was closed and because poaching and recreational fishing are highly unlikely given the remoteness and regular monitoring of the NWHI, tagged lobsters were only recaptured during the annual tag/recapture cruises. To prevent horizontal displacement of a lobster released at the surface from the effects of currents, all lobsters were released on the seafloor in the immediate area of capture (midpoint of the string of traps) via a release cage (O’Malley 2008). The ITP tag number, carapace length (CL; nearest 0.01 mm), sex, date of capture, and position of release (lati- tude and longitude determined using a Global Positioning System unit) were recorded for each tagged and recaptured lobster. Carapace length was measured as the straight-line distance between the rostrum and the posterior edge of the carapace along the dorsal midline. Each tag/recapture cruise had at least one tagger from previous cruises, which provided continuity in the tagging process. Data Analysis.—Distance moved (km) between release and recapture locations was calculated using spherical trigonometry (i.e., Great Circle Distance; Beyer and Shelby 1976). All positions were plotted using ArcGIS and visually validated; obviously erroneous release or recapture positions (e.g., on land or in locations where tagging operations did not take place) were removed. Distance moved was log-transformed to meet assumptions of normality. The effects of species, bank, sex, and CL-at-release on distance moved were tested by analyses of covariance (ANCOVA). The Rayleigh z-test of circular distribution (Zar 1999) was used to test the null hypothesis that the direction of lobster movement was random with respect to the release position (i.e., that the direction of population movement was uniformly distributed around a circle). Rejection of the null hypothesis (P < 0.05) would indicate a mean population movement direction. To determine if lobsters exhibited migrations toward or away from deeper-water habitat [20 fathom (37 m) contour], the distance and angle from the release position to the 20 fathom contour were determined using the ArcGIS 9.3 NEAR tool. Two temporal comparisons of lobster movement patterns were made. First, the timing of the annual tagging cruises allowed the recapture data to be partitioned into an annual (approximately 365 d) data set allowing annual movement patterns to be directly assessed. Second, long-term movements were determined by examining the data set in its entirety (individuals at liberty >1 yr). If individuals were recaptured more than once, only the initial and final capture data were used in the analysis. This was done to ensure equal weight in the analysis of any individual’s specific movement behavior and to maximize the time at liberty. No immediate recaptures (i.e., lobsters captured during the same tagging cruise) were included in either data set.

Results

Tagging and Recaptures.—In total, 54,528 P. marginatus and 30,218 S. squammosus were tagged from 2003 through 2008 at all locations combined. Bank-specific recapture rates ranged from 3.8% to 10.8% and from 9.6% to 11.6% for P. marginatus and S. squammosus, respectively. Male P. marginatus CL-at- tagging ranged from 42.0 to 133.3 mm [mean 86.1 (SD 12.6) and females ranged from 38.4 to 115.3 mm (mean 82.9 (SD 11.0)]. Male S. squammosus CL-at-tagging ranged from 55.1 to 94.6 mm [mean 75.9 (SD 5.3); female CL ranged from 56.1 to 102.8 mm (mean 78.9 (SD 6.3)]. o’malley and walsh: NWHI lobster movements 535

Between-Bank Movement.—No tagged individuals of either species moved between banks. All individuals were recaptured at the same bank at which they were tagged. Within-Bank Annual Movement.—Data from 2531 P. marginatus that averaged 354 days at liberty (DAL; SD 28) and 2119 S. squammosus that averaged 358 DAL (SD 30) before recapture were used for the analysis of annual movements. A preliminary ANCOVA indicated that there were significant interspecific differences in the annual distances moved by P. marginatus and S. squammosus (F1,4649 = 12.14, P < 0.001). Therefore, subsequent analyses were conducted at a species- specific level. Annual distances moved by P. marginatus varied significantly among banks

(ANCOVA: F 3,2525 = 14.29, P < 0.0001) and between sexes (ANCOVA: F1,2525 = 4.76, P = 0.03; Fig. 3). The size of P. marginatus did not affect distances moved

(ANCOVA: F 1,2525 = 2.14, P = 0.14). Despite the statistical differences among banks and between sexes, the large majority (97%) of male and female P. marginatus at all banks moved <1 km after 1 year at liberty (YAL). A small minority of P. marginatus (0.5%) moved at least 5 km (Fig. 3). Annual distances moved by S. squammosus also varied among banks (ANCOVA:

F3,2113 = 11.54, P < 0.0001) and between sexes (ANCOVA: F1,2113 = 4.09, P = 0.04;

Fig. 4). The size of S. squammosus had a significant positive effect (ANCOVA: F1,2113 = 7.56, P = 0.006) on distances moved with large individuals moving greater distances. Similar to P. marginatus and, despite the significant differences among banks, between sexes, and by size, the large majority (94%) of male and female S. squammosus at all banks moved <1 km annually. Two percent of tagged S. squammosus moved 5 km or more (Fig. 4). Within-Bank Long-Term Movement.—Movements of 4170 P. marginatus [DAL = 572 (SD 338)] and 3215 S. squammosus [DAL = 537 (SD 302)] were used to investigate whether there were long-term (>1 YAL) movement patterns. A preliminary ANCOVA demonstrated that there were significant interspecific differences in the distances moved (F1,7366 = 21.50, P < 0.0001); therefore, subsequent analyses were performed at a species-specific level. Long-term lobster movement patterns were very similar to annual movement patterns in most respects. Panulirus marginatus movements were affected by bank

(ANCOVA: F 3,4151 = 6.48, P = 0.0002); however, there was no apparent pattern.

Distance moved varied by sex (ANCOVA: F1,4151 = 5.33, P = 0.02), but the only discernible pattern was Necker Island females moving greater distances than males regardless of YAL. YAL also affected distance moved (ANCOVA: F4,4151 = 8.06, P < 0.0001), but with no discernible pattern. Several interactions were significant, including those between size and bank (ANCOVA: F3,4151 = 5.56, P = 0.0008), size and sex (ANCOVA: F1,4151 = 6.96, P = 0.008), and bank and YAL (ANCOVA:

F5,4151 = 5.91, P < 0.0001), indicating variable influences of these factors on distances moved. This is likely the reason why patterns are not apparent within any of these individual factors. Mean distances moved by P. marginatus at liberty for 2, 3, 4, and 5 yrs were <1 km (Fig. 3). The two exceptions were (1) males at Gardner Pinnacles at liberty for 2 yrs, altough this was likely a function of the small sample size and one individual moving >5 km, and (2) females at Maro Reef at liberty for 536 BULLETIN OF MARINE SCIENCE. VOL 89, NO 2. 2013 male (dark bars) and female (open bars) mean (SD) distance each year for mean at liberty Necker malebars) Island, Gardner moved and bars) (SD) (dark female (B) (open at (A) Panulirus marginatus Panulirus Figure 3. Pinnacles, error Maro above and Laysan (C) bars Reef, Island. (D) Values are sample sizes. Circles percent indicate recaptures of and male female (open) (dark) km.that <1 moved o’malley and walsh: NWHI lobster movements 537 male (dark bars) and female (open bars) mean (SD) distance each year for mean at liberty male bars) Necker and moved Island, Gardner bars) (SD) (dark female (open (B) at (A) Figure Scyllarides 4. squammosus Pinnacles, error Maro above and Laysan (C) bars Reef, Island. (D) Values are sample sizes. Circles percent indicate recaptures of and male female (open) (dark) km.that <1 moved 538 BULLETIN OF MARINE SCIENCE. VOL 89, NO 2. 2013

3 yrs, but this involved only one recapture. Ninety-seven percent of lobsters at liberty for >1 yr moved <1 km (Fig. 3) and <1% moved >5 km. Long-term movements of S. squammosus also differed significantly among banks

(ANCOVA: F 3,3200 = 35.73, P < 0.0001), but there were no detectable patterns (Fig.

4). Long-term movement varied between sexes (ANCOVA: F1,3200 = 3.9, P = 0.04) with, in most cases, females moving greater distances than males (Fig. 4). YAL also affected distance moved (ANCOVA: F4,3200 = 9.27, P < 0.001) but there were no patterns. There was also a significant size effect (ANCOVA: F1,3200 = 4.95, P = 0.03) with larger individuals moving greater distances. The interaction of banks and YAL (ANCOVA: F5,3200 = 7.63, P < 0.0001) also was significant, contributing to the lack of apparent patterns within any of these individual factors. The majority of S. squammosus at liberty for 2, 3, 4, and 5 yrs moved very little, <1 km on average (Fig. 4). The exceptions were Maro Reef males at liberty for 2 yrs (1.10 km) and Maro Reef females at liberty for 4 yrs (1.43 km). Ninety-two percent of S. squammosus at liberty for >1 yr moved <1 km and 2.8% moved 5 km or more (Fig. 4). Multiple Individual Recaptures.—Eleven Necker Island P. marginatus, and two Necker Island and 10 Maro Reef S. squammosus were recaptured in at least 3 consecutive yrs. Of these, only one P. marginatus and one S. squammosus moved >1 km from the initial tagging location (Table 1). Lobsters of both species tended to move within a specific home range; there was no migration in a specific direction away from the original release site (Figs. 5, 6). Point to Edge.—Proximity to the deep water drop-off did not affect the distances moved by P. marginatus of either sex at any bank (ANOVA: all Ps > 0.05). The same was true for S. squammosus (at all islands; ANOVA: P > 0.05) except at Maro Reef, where both males (ANOVA: F1,851 = 19.45, P < 0.0001) and females

(ANOVA: F 1,497 = 30.38, P < 0.0001) that were tagged farther away from the drop- off moved greater distances than those tagged near the drop-offF ( ig. 7). Individuals of both lobster species that were tagged and released at depths less than the 20 fathom contour did not appear to move toward deep water, and individuals that were tagged and released at depths greater than the 20 fathom contour did not appear to move toward shallower water. On average, 24% of lobsters moved within a 90° band in the direction of the 20 fathom contour (Fig. 8), while the random expectation would be 25%. Even when the analysis was restricted to individuals that moved more than 2 and 5 km, lobsters did not exhibit directed movements toward or away from deep water (Fig. 8). Vector Analysis.—Direction of travel was uniformly distributed for 11 of the species/sex/bank data sets (Table 2). However, significant departures from ran- domness were observed in five data sets: male P. marginatus at Necker Island and Maro Reef, male S. squammosus at Gardner Pinnacles and Maro Reef, and female S. squammosus at Maro Reef (Table 2). The mean direction of movement for these lobsters was between west and south-southwest. o’malley and walsh: NWHI lobster movements 539 0.03 recap 4 km from 5 YAL 0.11 tagging km from 0.03 0.02 0.02 0.02 0.07 recap 3 km from 4 YAL 0.12 0.13 0.05 0.13 0.09 tagging km from 0.22 0.02 0.01 0.02 0.02 0.29 0.34 0.04 0.02 0.03 0.03 0.13 0.03 0.10 0.01 0.12 0.04 0.45 0.45 0.12 0.02 0.03 1.53 recap 2 km from 3 YAL 0.11 0.11 1.07 0.03 0.31 0.44 0.14 0.30 0.19 0.05 0.14 0.03 0.00 0.06 0.08 0.04 0.04 0.44 0.44 0.04 0.08 0.03 0.14 tagging km from 0.81 0.03 0.06 0.04 0.03 0.01 0.36 0.08 0.03 0.10 0.04 0.10 0.02 0.04 0.07 0.04 0.02 0.02 0.02 0.06 0.17 0.05 1.72 recap 1 km from distance (km) moved from original tagging location and from previous recapture location for and from previous recapture location tagging location (km) moved from original distance 2 YAL 1.10 0.01 0.32 0.45 0.13 0.01 0.34 0.02 0.13 0.08 0.03 0.08 0.14 0.08 0.08 0.08 0.05 0.01 0.01 0.13 0.10 0.04 1.63 tagging km from 0.28 0.04 0.27 0.47 0.10 0.01 0.02 0.07 0.10 0.03 0.02 0.03 0.15 0.07 0.16 0.06 0.04 0.02 0.02 0.18 0.18 0.04 0.13 1 YAL tagging km from F F F F F F F F F F F F M M M M M M M M M M M Sex Maro Bank Maro Maro Maro Maro Maro Maro Maro Maro Maro Necker Necker Necker Necker Necker Necker Necker Necker Necker Necker Necker Necker Necker S. squammosus Scyllarides squammosus and Scyllarides 1. Panulirus marginatus Table = years at liberty. YAL individuals at liberty for >3 consecutive years. Species P. marginatus P. 540 BULLETIN OF MARINE SCIENCE. VOL 89, NO 2. 2013

Figure 5. Individual Panulirus marginatus movement patterns determined from consecutive annual (month/year) recaptures. Note different scales for the distance moved.

Figure 6. Individual Scyllarides squammosus movement patterns determined from consecutive annual (month/year) recaptures. Note different scales for the distance moved. o’malley and walsh: NWHI lobster movements 541

Figure 7. Relationship between distance to edge (20 fathom contour) and distance moved for male (dark circles) and female (open squares) Scyllarides squammosus at Maro Reef.

Discussion

Tag/Recapture Assumptions.—Despite the extensive use of mark/recapture data to examine lobster movements, Herrnkind (1980) identified three specific issues that need to be addressed in relation to the present study. The first, that probability dictates that most tagged animals will be recaptured soon after release near the release point, was not true of this study because all recaptures occurred after approximately 1 YAL. This gave the lobsters’ ample time to adjust to the tagging procedure and resume normal movement behavior. The second, that the greatest proportion of the recaptures will occur where fishing is heaviest, even if the animals disperse randomly, also did not apply to our study because the fishery was closed during the study period and all recaptures occurred during tag/recapture cruises. Further, the survey design ensured that banks were fished in their entirety when possible, increasing the probability of recapturing individuals that moved greater distances or to different habitats. Herrnkind’s (1980) final concern was the induction of nomadism by displacement of lobsters from the home reef during the tagging process. This effect was minimized in the present study by releasing lobsters on the seafloor as close as possible to the specific area of capture (within 366 m). Lack of NWHI Lobster Movement.—This study found that lobsters do not move between banks within the NWHI, which is consistent with the findings of Shaklee and Samollow (1984) and Seeb et al. (1990) that NWHI lobster populations are connected only by larval dispersal. This study also revealed that the vast majority of NWHI P. marginatus and S. squammosus moved <1 km from the release site, which corresponded to Herrnkind’s (1980) category of “homing-ter- ritorial.” Multiple consecutive annual recaptures of individuals confirm this by showing that NWHI lobsters move within a home range. Although a very small 542 BULLETIN OF MARINE SCIENCE. VOL 89, NO 2. 2013

Figure 8. Percent of male and female spiny lobster, Panulirus marginatus, and slipper lobster, Scyllarides squammosus, that traveled within a 90° angle towards the shelf edge (20 fathom contour). Results are plotted as lobsters that moved all distances (open bars), >2 km (gray bars), and >5 km (black bars). percentage moved farther than 5 km, there is no evidence that either species regu- larly exhibit either of the two longer-distance movement behaviors of “nomadism” or “migration” (Booth 1997). Even after 5 YAL, the mean distance moved from the original tagging location was 0.09 km and 0.23 km for male and female P. marginatus, respectively, and 0.54 km and 0.35 km for male and female S. squammosus, respectively. Sex was a significant factor in distances moved for both species, but the only obvious pattern was that female S. squammosus moved greater distances than male S. squammosus. This pattern was only true for 1 and 2 YAL; afterward, the consistency of females moving greater distances than males broke down. Size- at-tagging was a significant factor for only S. squammosus, for which distances moved increased with increasing CL. The magnitude of these size-specific movements is not necessarily indicative of large-scale unidirectional movements and may simply mean that larger S. squammosus have larger home ranges. Lobster tagging operations were confined to the June–September period, so seasonality of movements could not be assessed directly. But two pieces of cir- cumstantial evidence suggest that seasonal migrations to deep-water refuges [e.g., P. argus (Kanciruk and Herrnkind 1978)] or to access offshore currents [e.g., I. chacei (Stewart and Kennelly 1998)] did not take place. If there were seasonal movements toward deeper water, one would expect a greater possibility of individuals farther away from the deep water drop-off not homing to their exact home reef upon returning to the interior of the bank. This would be evident in the tagging data as individuals in the interior of the bank moving greater distances from the release site than individuals closer to the deep-water drop off. This was only true of male and female S. squammosus at Maro Reef; no other S. squammosus populations and none of the P. marginatus populations displayed this behavior. Second, it appears that individuals were not moving in the general direction of deep water. o’malley and walsh: NWHI lobster movements 543

Table 2. Rayleigh’s z-test results and mean direction moved for Panulirus marginatus and Scullarides squammosus.

Mean direction Species Sex Bank N z P moved (°) P. marginatus Male Necker Island 1,002 23.41 0.001 265.30 Gardner Pinnacles 30 2.69 0.10 > P > 0.05 Maro Reef 150 7.68 0.001 228.77 Laysan Island 9 0.38 > 0.50 Female Necker Island 1,062 1.65 > 0.20 Gardner Pinnacles 54 0.77 0.50 > P > 0.20 Maro Reef 209 2.25 0.20 > P > 0.10 Laysan Island 15 1.52 0.50 > P > 0.2 S. squammosus Male Necker Island 175 0.31 > 0.5 Gardner Pinnacles 200 6.55 0.0002 > P > 0.001 204.01 Maro Reef 854 3.21 0.05 > P > 0.002 238.05 Laysan Island 112 1.66 0.20 > P > 0.10 Female Necker Island 99 1.56 0.50 > P > 0.20 Gardner Pinnacles 94 0.41 0.50 > P > 0.20 Maro Reef 500 7.12 < 0.0001 256.47 Laysan Island 85 2.14 0.20 > P > 0.10

Even among the group of individuals that moved >5 km, <22% moved toward the deep-water drop off. A confounding situation arises from the few species-, sex-, and location-specific data sets displaying directed population movements (P. marginatus males at Necker Island and Gardner Pinnacles, and S. squammosus males at Gardner Pinnacles and Maro Reef and females at Maro Reef), but this is likely artifactual because there are no prominent features in the direction they moved. However, sampling was not conducted year-round so the seasonality of NWHI lobster movements, based on these data, remains unknown. Why Do NWHI Lobsters Not Move Much?—Why species that are evolutionarily and morphologically similar exhibit different movement patterns is currently unknown. Panulirus marginatus belongs to the group of Indo-West Pacific Panulirus species (Ptacek et al. 2001), which includes some species that exhibit on- togenetic movements, such as P. argus (Herrnkind 1980), Panulirus cygnus (Phillips 1983), and Panulirus japonicus (Yoshimura and Yamakawa 1988). The movement patterns of other species in the group, Panulirus femoristriga, Panulirus longipes, and Panulirus pascuensis, are currently unknown. However, other species within the family Palinuridae show high site fidelity. For instance, Galápagos Panulirus penicillatus (Hearn and Murillo 2008) and South African Panulirus homarus rubellus (Steyn and Schleyer 2011) do not exhibit large movements at any point in their life. Australian Panulirus versicolor also rarely move, with 55% found at or close to the den in which they were tagged; however, the limited spatial scale of the study may have biased estimates of distances moved (Frisch 2007). Only Panulirus guttatus in Florida, which spend their entire benthic lives on small portions of reefs (Sharp et al. 1997), are more restricted in their movement than the previously mentioned palinurids. There are few studies of scyllarid movements, but they document occurrenc- es of each of Herrnkind’s (1980) movement pattern categories. Some move very 544 BULLETIN OF MARINE SCIENCE. VOL 89, NO 2. 2013 little (i.e., homing), such as Australian Ibacus peronii (Stewart and Kennelly 1998), and Galápagos Scyllarides astori (Hearn 2006). Scyllarides aequinoctialis in Florida also display a high degree of site fidelity, but there is some speculation that ovigerous females may move to deeper waters to spawn (Sharp et al. 2007). Two scyllarid species fit the longer distance categories of nomadism and migration. Australian Thennus sp. moved a mean of 24 km, but there is no directed movement; thence, they are considered nomads (Jones 2007). They prefer featureless substrate, which provides no shelter to repetitively locate. Australian east coast Ibacus chacei display contranatant migrations, with adults migrating an average of 28–45 km northward to spawn where the East Australian Current transports the larvae southward to juvenile habitat (Stewart and Kennelly 1998). Stewart and Kennelly (1998) remark that Ibacus peronii and Ibacus chacei are morphologically nearly identical and overlap in their distributions, yet display different patterns of movement. Movement patterns within a given scyllarid species can also vary spatially. Mediterranean Sea Scyllarides latus in the Strait of Sicily move very little (Bianchini et al. 2001), while those off the coast of Israel move to deeper waters in August/ September (Spanier et al. 1988). Lavalli et al. (2007) speculated that water temperature is the driving force with the Israel population moving to deeper, cooler water to meet the physiological and behavioral requirements for molting, whereas the waters in the Strait of Sicily do not become as seasonally warm, thereby elimi- nating the necessity for these lobsters to move to molt. Childress and Jury (2006) summarized the most common explanations for directed movement as (1) enhancement of growth via increased degree-days or feeding opportunities, (2) facilitation of offspring release, and (3) avoidance of suboptimal habitats. Movement of juvenile NWHI P. marginatus and S. squammosus to adult habitat is unlikely due to the lack of typical lobster nursery habitats [e.g., mangroves, seagrass (Butler et al. 2006)] in the NWHI (Friedlander et al. 2009), which is consistent with the hypothesis of Parrish and Polovina (1994) that puerulus settlement occurs to adult habitat. Due to the small size of the banks, larval access to offshore currents is likely not important (Parrish and Polovina 1994); thus, adult females do not need to move toward the edge of the banks to release their eggs (e.g., P. cygnus, I. chacei). Finally, although the NWHI undergo a seasonal shift in oceanographic conditions (Desch et al. 2009), there is no large change in water temperature such as that which triggers the northern Caribbean and Bahamas P. argus mass offshore migration (Kanciruk and Herrnkind 1978). NWHI lobsters may not move for the simple reasons that there is no place to move to, the juvenile and adult habitats are the same, and offshore currents are within reach of newly hatched larvae. NWHI Ecosystem and Management Implications.—Our results shed light on spatial recruitment processes at individual banks within the NWHI. Both lobster species studied are characterized by long pelagic larval durations, approximately 1 yr for P. marginatus and 6–9 mo for S. squammosus, during which they are transported at least 25 nmi offshore (Polovina and Moffitt 1995). The lack of movement of juveniles and adults suggests that once late stage phyllosoma are near the NWHI they may be entrained in eddies or currents, which deliver them en masse to specific locations within the bank rather than distribute them evenly o’malley and walsh: NWHI lobster movements 545 across the bank. Panulirus marginatus puerulus and S. squammosus nisto settle, metamorphose, and probably do not move extensively because (1) juvenile and adult habitats are the same and (2) migrations across the bank would expose them to high levels of predation. Our results also explain the differences in size-frequency distributions within Necker Island. Tagging data did not indicate size-specific migrations of P. marginatus or S. squammosus, which differs from previous assumptions that lobster nursery habitats exist in the NWHI (DiNardo and Marshall 2001). Uneven settlement of P. marginatus puerulus across Necker Island may have resulted in relatively more individuals in the northern portion, the previously assumed nursery area. The commercial fishery targeted this area because of greater lobster abundance, which resulted in the removal of larger individuals from that area. This combina- tion of large numbers of small individuals from high levels of recruitment plus removal of large individuals from high levels of fishing effort is the most plausible explanation for spatial variability in the size-frequency distribution across Necker Island that was interpreted as an indicator of the existence of a nursery area. Species with a sedentary lifestyle or that move very little are highly susceptible to localized depletion. Areas that experience intense fishing effort must then be replenished by post-larval recruitment. Given the potentially patchy nature of lobster larval settlement across a given NWHI bank and the small home ranges of juveniles and adults of these species, heavily fished areas might not recover from exploitation for extended periods that cannot be predicted. Localized depletion of lobsters may have wide-ranging impacts to the marine ecosystems via fishery- induced trophic cascades (Salomon et al. 2010). Spatial management, in particular marine protected areas (MPA), is a technical measure that increasingly is becoming part of a comprehensive management plan for exploited species (Lorenzen et al. 2010). MPAs offer benefits to fisheries via two mechanisms, spillover from the protected area to the adjacent fishing area and increased egg production due to increased biomass and abundance within the MPA (Ward et al. 2001). The effectiveness of MPAs depends on the size of the MPA, the habitat within and bounding the MPA, and how these relate to the movement patterns of the species of interest (Moffitt et al. 2011, Moland et al. 2011). For instance, a MPA that is small relative to a species home range and is bounded by habitat that does not constrain movements across the boundary will fail to protect species because of excessive spillover to the adjacent fishing areas. That being said, if they are well designed F( reeman et al. 2009), even relatively small MPAs can benefit species that display high site fidelity such as P. marginatus and S. squammosus, and thereby their associated fisheries, through increased egg production and larval export (Diaz et al. 2011). Several studies have shown that relatively small MPAs (1–4 km2) are effective at retaining lobsters that show high site fidelity including Palinurus elephas (Goñi 2006, Follesa et al. 2007), Jasus edwardsii (Barrett et al. 2009), and Homarus americanus (Rowe 2001). The system of MPAs in place in the main Hawaiian Islands generally is considered too small and haphazardly designed to provide adequate species protection and any fisheries benefits, especially for highly mobile species F( riedlander et al. 2007). However, how effective they are at protecting lobsters while enhancing production outside the MPAs is currently unknown. The exact benefits can only be assessed by -ex amining the distance moved by these species in relation to the size of and the 546 BULLETIN OF MARINE SCIENCE. VOL 89, NO 2. 2013 amount of suitable habitat within each MPA, as well as determining the habitat that defines an individual’s home range. In conclusion, based on our movement analyses, NWHI P. marginatus and S. squammosus do not move appreciable distances, even after spending long times at liberty, and appear to exhibit high site fidelity. However, both species exhibited the capacity to move greater distances, as evidenced by a small minority of individuals traveling up to several kilometers. There was minimal evidence that the movements were directed, seasonal, or size-based migrations. Lobster movement patterns cannot be considered an evolutionarily conserved behavior because congeners of NHWI P. marginatus and S. squammosus exhibit movements that range from sedentary to migratory. Therefore, lobster movement patterns, including those we described, are likely driven by characteristics of local habitats. The lack of movements also suggests that small, yet well-designed MPAs could greatly enhance benefits to these species and their fisheries.

Acknowledgments

We thank the scientists who participated in the tagging cruises and the captain and crew of the F/V Marie M and F/V Katy Mary. We also thank T Rippetoe for GIS assistance. R O’Conner and T Acoba provided bathymetry maps. Helpful comments were provided in re- views by M Iacchei, M Musyl, and three anonymous reviewers. G DiNardo secured financial support via the National Oceanic and Atmospheric Administration Cooperative Research Program. Northwestern Hawaiian Islands activity was conducted under permit numbers NWHICRER-2006-003, PMNM-2007-023, and PMNM-2008-044.

Literature Cited

Barrett N, Buxton C, Gardner C. 2009. Rock lobster movement patterns and population structure within a Tasmanian marine protected area inform fishery and conservation management. Mar Freshwat Res. 60:417–425. http://dx.doi.org/10.1071/MF07154 Beyer WH, Selby SM. 1976. CRC Standard Mathematical Tables, 24th edition. CRC Press, Cleveland, OH. Bianchini ML, Bono G, Ragonese S. 2001. Long-term recaptures and growth of slipper lobsters, Scyllarides latus, in the Strait of Sicily (Mediterranean Sea). Crustaceana. 74:673– 680. http://dx.doi.org/10.1163/156854001750377948 Booth JD. 1997. Long-distance movements in Jasus spp. and their role in larval recruitment. Bull Mar Sci. 61:111–128. Butler MJ IV, Dolan TW III, Hunt JH, Rose KA, Herrnkind WF. 2005. Recruitment in degraded marine habitats: a spatially explicit, individual-based model for spiny lobster. Ecol Appl. 15:902–918. http://dx.doi.org/10.1890/04-1081 Butler MJ, Steneck RS, Hernkind WF. 2006. Juvenile and adult ecology. In: Phillips BF, editor. Lobsters: biology, management, aquaculture, and fisheries. Blackwell Scientific Publications, Oxford. p. 263–309. PMCid:1779709. http://dx.doi. org/10.1002/9780470995969.ch8 Childress MJ, Jury SH. 2006. Behavior. In: Phillips BF, editor. Lobsters: biology, management, aquaculture, and fisheries. Blackwell Scientific Publications, Oxford. p. 78–112. http://dx.doi.org/10.1002/9780470995969.ch3 Davis GE, Dodrill JW. 1989. Recreational fishery and population dynamics of spiny lobsters, Panulirus argus, in Florida Bay, Everglades National Park, 1977–1980. Bull Mar Sci. 44:78–88. o’malley and walsh: NWHI lobster movements 547

Desch A, Wynne T, Brainard R, Friedlander A, Christensen J. 2009. Oceanographic and physical setting. In: Friedlander A, Keller K, Wedding L, Clarke A, Monaco M, editors. A marine biogeographic assessment of the Northwestern Hawaiian Islands. NOAA Tech. Memo. NOS NCCOS 84, Silver Springs. p. 17–63. Díaz D, Mallol S, Parma AM, Goñi R. 2011. Decadal trend in lobster reproductive output from a temperate marine protected area. Mar Ecol Prog Ser. 433:149–157. http://dx.doi. org/10.3354/meps09182 DiNardo GT, Marshall R. 2001. Status of lobster stocks in the Northwestern Hawaiian Islands, 1998–2000. Honolulu Lab, Southwest Fish Sci Cent, Natl Mar Fish Serv, NOAA, Honolulu, HI 96822-2396. Southwest Fish Sci Cent Admin Rep. H-01-04. 47 p. Follesa CM, Cuccu D, Cannas R, Sabatini A, Cau A. 2007. Emigration and retention of Palinurus elephas (Fabricius, 1789) in a central western Mediterranean marine protected area. Sci Mar. 71:279–285. Freeman DJ, MacDiarmid AB, Taylor RB. 2009. Habitat patches that cross marine reserve boundaries: consequences for the lobster Jasus edwardsii. Mar Ecol Prog Ser. 388:159– 167. http://dx.doi.org/10.3354/meps08122 Friedlander AM, Brown EK, Monaco ME. 2007. Coupling ecology and GIS to evaluate ef- ficacy of marine protected areas in Hawaii. Ecol Appl. 17:715–730. PMid:17494391. http://dx.doi.org/10.1890/06-0536 Friedlander A, Keller K, Wedding L, Clarke A, Monaco M. 2009. A marine biogeographic assessment of the Northwestern Hawaiian Islands. NOAA Tech Memo. NOS NCCOS 84, Silver Springs. 363 p. Frisch AJ. 2007. Short- and long-term movements of painted lobster (Panulirus versicolor) on a coral reef at Northwest Island, Australia. Coral Reefs. 26:311–317. http://dx.doi. org/10.1007/s00338-006-0194-6 Goethel DR, Quinn TJ, Cadrin SX. 2011. Incorporating spatial structure in stock assessment: movement modeling in marine fish population dynamics. Rev Fish Sci. 19:119– 136. http://dx.doi.org/10.1080/10641262.2011.557451 Goñi R, Quetglas A, Renones O. 2006. Spillover of spiny lobsters Palinurus elephas from a marine reserve to an adjoining fishery. Mar Ecol Prog Ser. 308:207–219. http://dx.doi. org/10.3354/meps308207 Hearn A. 2006. Life history of the slipper lobsters Scyllarides astori Holthuis 1960, in the Galápagos Islands, Ecuador. J Exp Mar Biol Ecol. 328:87–97. http://dx.doi.org/10.1016/j. jembe.2005.06.021 Hearn A, Murillo JC. 2008. Life history of the red spiny lobster, Panulirus penicillatus (Decapoda: Palinuridae), in the Galápagos Marine Reserve, Ecuador. Pac Sci. 62:191– 204. http://dx.doi.org/10.2984/1534-6188(2008)62[191:LHOTRS]2.0.CO;2 Herrnkind WF. 1980. Spiny lobsters: patterns of movement. In: Cobb JS, Phillips BF, editors. The biology and management of lobsters, Vol 1. Physiology and behaviour. Academic Press, New York. p. 349–407. Jones CM. 2007. Biology and fishery of the bay lobster,Thenus spp. In: Lavalli KL, Spanier E, editors. The biology and fisheries of the slipper lobster. CRC, Boca Raton. p. 326–358. http://dx.doi.org/10.1201/9781420005165.ch16 Kanciruk P, Herrnkind WF. 1978. Mass migration of spiny lobster, Panulirus argus (Crustacea: Palinuridae): behavior and environmental correlates. Bull Mar Sci. 28:601–623. Lavalli K, Spanier E, Grasso F. 2007. Behavior and sensory biology of slipper lobsters. In: Lavalli KL, Spanier E, editors. The biology and fisheries of the slipper lobster. CRC, Boca Raton. p. 133–181. http://dx.doi.org/10.1201/9781420005165.ch7 Lorenzen K, Steneck RS, Warner RR, Parma AM, Coleman FC, Leber KM. 2010. The spatial dimensions of fisheries: putting it all in place. Bull Mar Sci. 86:169–177. MacDonald CD, Jazwinski SC, Prescott JH. 1984. Queuing behavior of the Hawaiian spiny lobster Panulirus marginatus. Bull Mar Sci. 35:111–114. 548 BULLETIN OF MARINE SCIENCE. VOL 89, NO 2. 2013

Moffitt EA, White JW, Botsford LW. 2011. The utility and limitations of size and spacing guidelines for designing marine protected area (MPA) networks. Biol Conserv. 144:306– 318. http://dx.doi.org/10.1016/j.biocon.2010.09.008 Moland E, Olsen EM, Andvord K, Knutsen JA, Stenseth NC. 2011. Home range of European lobster (Homarus gammarus) in a marine reserve: implications for future reserve design. Can J Fish Aquat Sci. 68:1197–1210. http://dx.doi.org/10.1139/f2011-053 O’Malley JM. 2008. Evaluations of tag retention and a device for releasing discarded Hawaiian spiny lobsters Panulirus marginatus. N Am J Fish Manage. 28:619–624. http:// dx.doi.org/10.1577/M07-102.1 Parrish FA, Polovina JP. 1994. Habitat thresholds and bottlenecks in production of the spiny lobster (Panulirus marginatus) in the Northwestern Hawaiian Islands. Bull Mar Sci. 54:151–163. Phillips BF. 1983. Migrations of pre-adult western rock lobster Panulirus cygnus, in Western Australia. Mar Biol. 76:311–318. http://dx.doi.org/10.1007/BF00393034 Polovina JJ, Moffitt RB. 1995. Spatial and temporal distribution of the phyllosoma of the spiny lobster, Panulirus marginatus, in the Northwestern Hawaiian Islands. Bull Mar Sci. 56:406–417. Ptacek MB, Sarver SK, Childress MJ, Herrnkind WF. 2001. Molecular phylogeny of the spiny lobster genus Panulirus (Decapoda: Palinuridae). Mar Freshwat Res. 52:1037–1047. http://dx.doi.org/10.1071/MF01070 Rowe S. 2001. Movement and harvesting mortality of American lobsters (Homarus americanus) tagged inside and outside no-take reserves in Bonavista Bay, Newfoundland. Can J Fish Aquat Sci. 58:1336–1346. http://dx.doi.org/10.1139/f01-083 Salomon AK, Gaichas SK, Shears NT, Smith JE, Madin EMP, Gaines SD. 2010. Key features and context-dependence of fishery-induced trophic cascades. Conserv Biol. 24:382–394. PMid:20151987. http://dx.doi.org/10.1111/j.1523-1739.2009.01436.x Seeb LW, Seeb JE, Polovina JJ. 1990. Genetic variation in highly exploited spiny lobster Panulirus marginatus populations from the Hawaiian Archipelago. Fish Bull. 88:713–718. Shaklee JB, Samollow PB. 1984. Genetic variation and population structure in a spiny lobster, Panulirus marginatus in the Hawaiian archipelago. Fish Bull. 82:693–702. Sharp WC, Hunt JH, Williams WG. 1997. Life history of the spotted spiny lobster, Panulirus guttatus, an obligate reef-dweller. Mar Freshwat Res. 48:687–698. http://dx.doi. org/10.1071/MF97125 Sharp WC, Hunt JH, Teehan WH. 2007. Observations on the ecology of Scyllarides aequinoctialis, Scyllarides nodifer, and Parribacus antarcticus and a description of the Florida Scyllarid fishery. In: Lavalli KL, Spanier E, editors. The biology and fisheries of the slipper lobster. CRC, Boca Raton. p. 231–242. PMid:17452528. PMCid:2064132. http:// dx.doi.org/10.1201/9781420005165.ch11 Spanier E, Tom M, Pisanty S, Almog G. 1988. Seasonality and shelter selection by the slipper lobster Scyllarides latus in the southeastern Mediterranean. Mar Ecol Prog Ser. 42:247– 255. http://dx.doi.org/10.3354/meps042247 Spanier E, Lavalli KL. 2006. Scyllarides species. In: Phillips BF, editor. Lobsters: biology, management, aquaculture, and fisheries. Blackwell Scientific Publications, Oxford. p. 462–496. http://dx.doi.org/10.1002/9780470995969.ch14 Stewart J, Kennelly SJ. 1998. Contrasting movements of two exploited Scyllarid lobsters of the genus Ibacus off the east coast of Australia. Fish Res. 36:127–132. http://dx.doi. org/10.1016/S0165-7836(98)00104-0 Steyn E, Schleyer MH. 2011. Movement patterns of the East Coast rock lobsters Panulirus homarus rubellus on the coast of KwaZulu-Natal, South Africa. N Z J Mar Freshwat Res. 45:85–101. http://dx.doi.org/10.1080/00288330.2010.538067 Uchida RN, Uchiyama JN, Tagami DT, Shiota PM. 1980. Biology, distribution, and estimates of apparent abundance of the spiny lobster, Panulirus marginatus (Quoy and Gaimard), in waters of the Northwestern Hawaiian Islands: Part 1. Distribution in relation to depth o’malley and walsh: NWHI lobster movements 549

and geographical areas and estimates of apparent abundance. In: Grigg RW, Pfund RT, editors. Proceedings of the symposium on status of resource investigations in the Northwestern Hawaiian Islands. Miscellaneous Report UNIHI-SEAGRANT-MR-80-04. Univ. of Hawaii Sea Grant College Program, Honolulu, HI. p. 121–130. Uchida RN, Uchiyama JN. 1986. Fishery atlas of the Northwestern Hawaiian Islands. US Dept of Commerce, NOAA Technical Report NMFS No. 38. 142 p. Ward TJ, Heinemann D, Evans N. 2001. The role of marine reserves as fisheries management tools: a review of concepts, evidence and international experience. Bureau of Rural Sciences, Canberra, Australia. 192 p. Yoshimura T, Yamakawa H. 1988. Ecological investigations of settled puerulus and juvenile stages of the Japanese spiny lobster Panulirus japonicus at Kominato, Japan. J Crust Biol. 8:524–531. http://dx.doi.org/10.2307/1548688 Zar JH. 1999. Biostatistical analysis, 4th ed. Prentice Hall, Upper Saddle River, NJ.

Date Submitted: 21 June, 2012. Date Accepted: 18 Janurary, 2013. Available Online: 26 February, 2013.

Addresses: (JMO) Joint Institute for Marine and Atmospheric Research, University of Hawai‘i 1000 Pope Street, Honolulu, Hawaii 96822. Email: . (WAW) Joint Institute for Marine and Atmospheric Research, University of Hawai‘i, 1000 Pope Street, Honolulu, Hawaii 96822. Email: .

Appendix 1. Species names and authorities in the present study. Panulirus argus (Latreille, 1804) Scyllarides latus (Latreille, 1802) Panulirus marginatus (Quoy and Gaimard, 1825) Scyllarides squammosus (H. Milne-Edwards, 1837) Scomber japonicus Houttuyn, 1782 Panulirus cygnus George, 1962 Panulirus japonicus (Von Siebold, 1824) Panulirus femoristriga Von Martens, 1872 Panulirus longipes (A. Milne-Edwards, 1868) Panulirus pascuensis Reed, 1954 Panulirus penicillatus (Olivier, 1791) Panulirus homarus rubellus Berry, 1974 Panulirus versicolor (Latreille, 1804) Panulirus guttatus (Latreille, 1804) Ibacus peronii Leach, 1815 Scyllarides astori Holthuis, 1960 Scyllarides aequinoctialis (Lund, 1793) Ibacus chacei Brown and Holthuis, 1998 Palinurus elephas (Fabricius, 1787) Jasus edwardsii (Hutton, 1875) Homarus americanus H. Milne-Edwards, 1837

&lt;I&gt;Panulirus Marginatus,&lt;/I&gt; and Slipper Lobster

<I>Panulirus Marginatus,</I> and Slipper Lobster