Alaska's Mandatory Shellfish Observer Program, 1988-2000

Crabs in Cold Water Regions: Biology, Management, and Economics 693 Alaska Sea Grant College Program • AK-SG-02-01, 2002

Alaska’s Mandatory Shellfish Observer Program, 1988-2000

Larry Boyle and Mary Schwenzfeier Alaska Department of Fish and Game, Division of Commercial Fisheries, Dutch Harbor, Alaska

Abstract The Alaska Department of Fish and Game (ADFG) initiated its Mandatory Shellfish Observer Program in September 1988. Industry concern over the department’s inability to monitor the harvest and processing of crabs by catcher-processor vessels and at-sea processor vessels prompted the Alaska Board of Fisheries (BOF) to implement the program. Guidelines for observed vessels, observer companies, and ADFG were established by the BOF. Changes enacted by the BOF over time addressed conflicts of interest, certification and decertification procedures for observers and observer companies, and increased observer qualifications. A federal research plan to fund state crab and federal groundfish observers through a 2% fee on landings was enacted under authority of the Magnuson Act in 1994, but was repealed in 1995. A department proposal to fund crab observer coverage from the harvest and sale of crabs was developed for industry review in 1996. The BOF adopted a modified proposal in 1999. Observer coverage has been implemented on catcher vessels in selected crab fisheries. Funds from harvest and sale of crabs pay for a portion of the observer costs. Current regulations allow the department to determine observer coverage levels to meet ADFG’s data collection needs. An industry task force appointed by the BOF advises ADFG and BOF on observer coverage and funding issues. While inception of the program was based on regulation enforcement, observers have become the primary means in most fisheries for gathering data for research, inseason management, and developing new management measures.

Introduction King and Tanner crab fisheries in the Bering Sea and Aleutian Islands (BSAI) are managed under a federal fisheries management plan (FMP). The FMP establishes a cooperative management regime that defers crab management 694 Boyle and Schwenzfeier — Alaska Shellfish Observer Program to the state of Alaska with federal oversight through the North Pacific Fishery Management Council (NPFMC 1998). The Alaska Board of Fisheries (BOF) is the regulatory body that establishes state fishery regulations within the framework of the FMP. The BOF authorized the Alaska Department of Fish and Game (ADFG) to implement its Mandatory Shellfish Observer Program in 1988. The department had traditionally collected essential biological and management data from crab catcher vessels at the point of landing, prior to processing. Industry evolution toward at-sea processing seriously eroded the department’s ability to adequately monitor harvests and collect essential biological data for inseason management. In 1987 catcher-processors made up 9% of the 236 vessels fishing red king crab (Paralithodes camtschaticus ) in the Bering Sea and accounted for 20% of the 12.3 million pound harvest (Morrison 1992). Department analysis of the 1987 Bristol Bay red king crab fishery detailed significant differences in catch rates between catcher vessels and catcher-processor vessels. Overall, catcher-processor vessels caught approximately 2.5 times more pounds of crab than catcher vessels. These differences were unre- lated to vessel length, number of pots fished, gear soak time, or areas fished (Schmidt and Johnson 1988). The department suspected that female crabs and undersized male red king crabs were being retained illegally and processed at sea by the catcher-processors. Without onboard observer coverage, female and undersized male crabs could be retained illegally and processed immediately, making it impossible to enforce size and sex regulations used to manage the fishery. During the first 2 years of the mandatory shellfish program, observers concentrated on sex and size regulation compliance in their duties aboard catcher-processors. The differences in catch and landing rates between catcher vessels and catcher-processors equalized after observers were placed on vessels in 1988. The mandatory onboard observer program has become an essential fisheries regulation and management tool. After the first 2 years of the program emphasis on data gathered has shifted away from regulation compliance to biological data gathering. The department and BOF now consult and use observer data for most BSAI shellfish management and regulatory decisions.

Initial Program In 1988 the BOF directed ADFG to institute a mandatory onboard shellfish observer program to promote both conservation and enforcement. The board found that:

• Onboard observers provide the only effective means of collecting essential biological and management data from catcher-processor and floating processor vessels that process shellfish at sea. Crabs in Cold Water Regions: Biology, Management, and Economics 695

• Fisheries onboard observers provide the only effective means to enforce regulations that protect shellfish resources.

• The cost of providing onboard observers is a reasonable expense to be borne by the processors.

• No acceptable management alternatives exist other than disallow- ing the operation of a vessel that refuses to cooperate with the onboard observer program.

The initial shellfish observer program required observers on all vessels that processed blue king crabs (P. platypus), golden king crabs (Lithodes aequispina), red king crabs, or Tanner crabs (Chionoecetes bairdi ) at sea in Alaska. Shellfish observers were first deployed on catcher-processor vessels during the 1988 Bristol Bay red king crab fishery. Observers were briefed and debriefed in the ADFG office responsible for managing those fisheries. Analysis of data from the 1988 Bristol Bay fishery found no significant difference in catch rates between catcher-processor vessels and catcher vessels in the fishery (Schmidt and Johnson 1989). Data from the 1989 Bristol Bay red king crab fishery yielded similar results, and Schmidt and Johnson (1990) concluded that the presence of fishery observers on the catcher-processor vessels contributed to the similarity in the catch rates. The following roles of the vessels, independent observer companies, ADFG, and observers were established by BOF.

Vessels Vessels contract and pay for observers through independent, third-party observer companies. While onboard, observers are provided food and accommodations equal to that of the vessel’s crew. Vessels must provide observers with a safe work area, totes to contain crab pot contents for sampling, and the opportunity to adequately sample the catch. Fishing effort, harvest, and location information is provided daily to observers. Access to communication equipment must also be provided for observers to communicate with the department for questions and information, and to transmit catch reports as specified by ADFG.

Observer Companies Observer companies recruit, hire, train, and deploy their at-sea employees and provide all administrative and payroll functions associated with their employment. They provide all logistical support for observers during training and deployment including food, accommodations, sampling equipment, safety gear, and transportation. Observer companies secure contracts for observer services directly with vessels. Currently, there are three observer providers certified by ADFG and active in the crab observer program. 696 Boyle and Schwenzfeier — Alaska Shellfish Observer Program

Alaska Department of Fish and Game The department is responsible for establishing policies and procedures for certification and decertification of contracting agents and observers. The department, for data consistency and reliability, developed observer training standards, sampling procedures, and sampling guidelines.

Observers An observer candidate must be sponsored by one of the observer companies and approved by the department. Current observer qualifications are a bachelor’s degree in biology or any branch of biology, or valid National Marine Fisheries Service (NMFS) observer certification. Observer candidates undergo training approved by ADFG and must pass a written exam. After successful completion of training, trainee observers attend a practical training exam administered by ADFG in Dutch Harbor. Trainee observers have 180 days to gain their certification by successfully completing assigned tasks during two, three, or four deployments depending on the individual observer’s data collection competence level. Certified observers who are inactive for 12 consecutive months lose their certification. Observers are limited to no more than 90 days on the same vessel during a 12 month period.

Revisions to the Shellfish Observer Program Subsequent revisions to the initial program have been made by the BOF. Mandatory observer coverage was expanded in 1991 to include all vessels that process snow crabs (C. opilio) at sea. The snow crab fishery was ex- cluded in the original mandatory observer program because the legal size for snow crabs of 3.1 inches in carapace width (cw) is substantially smaller than the market size of 4 inches cw. Processing of sublegal snow crabs had not been a problem; however, Bering Sea Tanner and snow crab stocks have overlapping ranges and the department had identified retention and processing of sublegal Tanner crabs as a problem during the snow crab fishery. Peak catcher-processor vessel participation and observer deployment days in the BSAI crab fisheries occurred in 1991 when 33 of these vessels were in operation (Table 1). Over time many of these vessels left U.S. fisheries for Russian waters as joint venture operations or were sold to Russian owners. In 1993 observer coverage was initiated for all vessels fishing under an ADFG commissioner’s permit in the Bering Sea for hair crab (Erimacrus isenbeckii ) or for any snail species because of management concerns about incidental bycatch of female and juvenile king crabs in these fisheries. Also during 1993, minimum qualifications for observers were upgraded to become uniform with the department’s fishery biologist level 1 job class qualifications. Current certification, decertification, and conflict of interest Crabs in Cold Water Regions: Biology, Management, and Economics 697

Table 1. Annual number of observer days of deployments by vessel type in the Bering Sea and Aleutian Islands shellfish fisheries, 1988- 1998.a

Catcher-processor Floating processor Catcher vessels vessels vessels Total

No. of Observer No. of Observer No. of Observer No. of Year vesselsb days vesselsb days vesselsb days observer days

1988 21 756 6 175 — — 931

1989 22 3,043 12 629 — — 3,672

1990 26 3,247 15 1,299 — — 4,546

1991 33 7,726 18 2,875 — — 10,601 1992 32 5,936 19 2,441 2 21 8,398 1993 29 3,670 21 1,766 14 853 6,289 1994 24 2,000 17 1,267 19 1,345 4,612 1995 21 1,728 15 843 50 3,477 6,048 1996 16 1,480 13 820 38 4,745 7,045 1997 15 1,635 11 882 30 2,356 4,873 1998 13 1,534 11 928 43 2,839 5,301 1999 10 1,120 11 850 41 1,893 3,863 2000 10 592 5 194 46 2,547 3,333 aExcludes scallop vessels. bUnique vessels.

guidelines were established for observer companies and observers by the BOF. The department also established a scallop observer program under the existing Shellfish Observer Program as part of an overall scallop management plan. An increase in the number of scallop vessels participating in Alaskan fisheries combined with lack of scallop stock data and crab bycatch numbers prompted the BOF to require 100% observer coverage on scallop vessels in most fishery management areas. A scallop FMP was developed in 1993 by the North Pacific Fishery Management Council (NPFMC) that allows nine scallop vessels to fish statewide (outside the Cook Inlet registration area). Crab observer coverage was implemented in 1994 on all vessels fishing under a commissioner’s permit for deepwater crab species in the Bering Sea, Aleutian Islands, South Peninsula, and Kodiak fishery registration areas. These species included grooved Tanner crab (C. tanneri ), triangle Tanner crab (C. angulatus), and scarlet king crab (Lithodes cousei ). Stock and life history information on these unexploited species are generally lacking. By 1994 a decrease in catcher-processor vessels participating in the Aleutian Islands king crab fisheries had severely reduced the department’s ability to collect data necessary for management of the fisheries. In 1994 698 Boyle and Schwenzfeier — Alaska Shellfish Observer Program the BOF adopted the department’s proposal to place observers on all vessels participating in the Aleutian Islands king crab fisheries. The coverage commenced at the start of the 1995 season.

North Pacific Fisheries Research Plan The NPFMC developed the North Pacific Fisheries Research Plan to fund both the NMFS North Pacific Groundfish Observer Program and the ADFG Shellfish Observer Program. The plan was to provide a framework for development and coordination of groundfish and crab observer programs and their effectiveness in meeting fishery management needs. This was authorized by the Magnuson Act passed in 1992. The council took final action on the plan in 1994. Fees of 2% were to be collected from the landings of groundfish and halibut harvested in the Exclusive Economic Zone (EEZ) off Alaska and all BSAI king and Tanner crabs. These fees would be used to fund the agency’s fishery observer programs. Fees collected in 1995 were scheduled to fund observer deployments in 1996. Observer companies were to bid on providing observers to NMFS and ADFG observer programs, which would deploy the observers. This would have reduced the conflict of interest inherent in the existing program by establishing an arms-length relationship between vessel companies and observer companies. Crab fisheries were assessed over $3.2 million dol- lars in 1995 and over $2.7 million of the fees were collected. However, the council repealed the plan later in 1995 after hearing concerns from portions of the crab and groundfish fishing industry on the structure of the fee system. Collected monies were refunded.

Proposal for a State-Funded Crab Observer Program After repeal of the North Pacific Fisheries Research Plan, ADFG proposed a plan in 1996 for a state-funded crab observer program for BSAI crab fisheries. The BOF had asked the department earlier that year to reevaluate the existing observer program. The proposal would fund all crab observer deployments from the harvest and sale of crabs by the department under its test-fishery receipt authority. Included in the proposal was an option for ADFG to hire and deploy seasonal employees as fishery observers. Previously, vessels that were required to carry observers absorbed additional costs compared to vessels not required to carry observers. Re- moving observer costs would eliminate the inequitable aspect of the program. Current costs for observer coverage are approximately $300 per day plus travel costs. Annual industry costs for observer coverage for the 1991 to 1998 period have ranged from an estimated $1.6 to $3.0 million, varying with season lengths and number of vessels participating. The number of catcher-processor vessels participating in BSAI fisheries had steadily decreased from 33 vessels in 1991 to only 10 by 1999 Crabs in Cold Water Regions: Biology, Management, and Economics 699

(Table 1). Collection of fishery data, based solely on catcher-processor vessel participation, was lacking or nonexistent in some of these valuable fisheries (Tables 2-4). Under existing regulations there was no provision for the department to require catcher vessels to carry observers in most of the Bering Sea king or Tanner crab fisheries. Data beneficial to management and research of these fisheries had become limited or nonexistent. The department’s proposal to fund crab observer deployments in the BSAI crab fisheries was deliberated by the BOF at their March 1999 meeting. However, the board and industry were not comfortable with funding the entire program from cost-recovery receipts. Revenue collection would pay for some observer deployments while remaining deployments would continue to be paid by the fishing vessels. New regulations passed by BOF gave the department authority to determine required observer coverage levels on catcher vessels to meet biological data gathering needs for each general fishery. Although the inception of the shellfish observer program was based on regulation enforcement, observers on crab fishing vessels have become the primary data gatherers for departmental research, inseason management, and the BOF. All vessels were required to preregister before participating in each fishery. This would enable the department to determine the number of observers to deploy based on the number of catcher vessels participating, and to use the preregistration list to randomly select catcher vessels to carry observers during the fishery. The BOF established an industry oversight task force to provide input to the department on observer coverage levels, funding, and other aspects of the crab observer program. To be effective, observer coverage levels must be flexible in each fishery depending on data needs and fishery characteristics. Deployment of crab observers under this program began on July 1, 2000.

Catcher Vessel Observer Coverage in Bering Sea Crab Fisheries The department collected cost-recovery receipts totaling $650,000 in 1999 to fund crab observer deployments from July 1, 2000 through June 30, 2001. This amount was less than 1% of the total 1999 ex-vessel value of the Bristol Bay red king crab fishery (Table 3). The oversight task force made several revisions to their original recommendations to the department as this new aspect of the program was developed. Their final recommendations were to use cost recovery funds only to pay for new observer coverage on catcher vessels. Vessels that previously paid for their observer coverage would continue to pay, and fisheries with 100% observer coverage would continue at that level. The task force’s recommendations were adopted for the first year. Crab observers would deploy on catcher vessels in the open access fisheries for Bering Sea king, Tanner, and snow crabs. Observer and associated 700 Boyle and Schwenzfeier — Alaska Shellfish Observer Program

Table 2. Historic vessel and observer participation, and ex- vessel values, St. Matthew and Pribilof king crab fisheries, 1989-1998.

Number of Percent Fishery Total observed catcher- of fleet ex-vessel value Year vessels processor vessels observed ($ millions)

St. Matthew king crab fishery 1989 69 15 21.7 3.5 1990 31 7 22.6 5.7 1991 68 9 13.2 9.0 1992 174 8 4.6 7.4 1993 92 3 3.3 9.7 1994 87 6 6.9 15.0 1995 90 1 1.1 7.1 1996 122 3 2.5 6.7 1997 117 1 0.9 9.8 1998a 131 2 2.3 5.3

Pribilof king crab fishery

1989 No fishery — — —

1990 No fishery — — — 1991 No fishery — — — 1992 No fishery — — — 1993 112 2 1.8 13.0 1994 104 0 0 8.6 1995 127 1 0.8 6.8 1996 66 0 0 3.0 1997 53 0 0 3.7 1998 55 0 0 2.4

a One catcher vessel also carried an observer during the fishery and is included in the percent of fleet observed. No fisheries occurred in 1999 or 2000. Crabs in Cold Water Regions: Biology, Management, and Economics 701

Table 3. Historic vessel and observer participation, and ex-vessel value, Bristol Bay red king crab fishery, 1988-2000.

Number of Number of Fishery observed observed ex-vessel Total catcher-processor catcher Percent of value Year vessels vessels vessels fleet observed ($ millions)

1988 200 20 0 10.0 37.6 1989 211 18 0 8.5 50.9 1990 240 20 0 8.3 101.2 1991 302 25 0 8.3 51.2 1992 281 18 0 6.4 40.2 1993 292 17 0 5.8 55.1

1994 No fishery — — — —

1995 No fishery — — — — 1996 196 4 0 2.0 33.6 a 1997 256 8 11 7.4 28.5 a 1998 275 11 10 7.6 37.4 1999 259 8 0 3.1 69.1 a 2000 245 7 14 8.6 36.0

aIncludes ADFG observers.

deployment costs would be paid for with the cost-recovery monies collected in 1999. The goals the first year were to deploy observers on 10% of the catcher vessels, with a minimum of five observers deployed on vessels 125 ft in length and minimum of five observers on vessels over 125 ft in length. Vessel pot limits in these FMP fisheries are based on the two vessel size categories. Vessels were selected for observer coverage at ran- dom from the preregistration list for each fishery. The first state-funded crab observer deployments on catcher vessels occurred during the Bristol Bay red king crab fishery in October 2000. There were 245 catcher vessels that preregistered for the fishery: 68 vessels were over 125 ft and 177 were 125 ft or less. Observers were deployed on five catcher vessels over 125 ft and six vessels 125 ft or less. The goal of 10% observer coverage on catcher vessels was not met due to a lack of available crab observers. Seven catcher-processors and one floating processor carried observers under the existing “pay-as-you-go” system. In addition, three catcher vessels fishing in a 25-vessel cooperative which was operating under federal American Fisheries Act regulations carried observers. The cooperative paid those observer costs. Overall, the 22 observers deployed at sea to collect fishery data accounted for nearly 9% of all vessels participating in the 2000 Bristol Bay red king crab fishery. 702 Boyle and Schwenzfeier — Alaska Shellfish Observer Program

Table 4. Historic vessel and observer participation, and ex-vessel values, Bering Sea snow crab and C. bairdi fisheries.

No. of Fishery ex-vessel Total observed catcher- Percent of value Fishery vessels processor vessels fleet observed ($ millions)

Bering Sea snow crab fishery, 1991-2000 1991 220 26 11.8 162.6 1992 250 30 12.0 156.5 1993 254 25 9.8 171.9 1994 273 24 8.8 192.4 1995 253 19 7.5 180.0 1996 234 15 6.4 85.6 1997 226 13 5.8 92.6 1998 229 12 5.2 134.6 1999 236 10 4.2 160.8 2000 231 8 3.5 56.3

Bering Sea C. bairdi directed fishery, 1991-2000 1991-1992 255 27 10.6 47.3 1992-1993 285 22 7.7 58.8 1993-1994 283 17 6.0 24.0 1994 183 9 4.9 28.5 1995 196 11 5.6 11.7 1996 135 2 1.5 2.0

1997-2000 No fishery — — —

Overview of the ADFG Shellfish Observer Program The ADFG Observer Program staff coordinate with observer companies to schedule crab observer briefings and debriefings in the ADFG office in Dutch Harbor. The observer program staff evaluates observer performance, certifies observers and observer companies, and tracks observer deployments. Observer documentation of potential fishing violations and any evidence collected are referred to the Alaska Department of Public Safety, Fish and Wildlife Protection Division, for processing. The observer program staff coordinates with ADFG shellfish research staff on observer sampling goals and methodologies. Catch, effort, bycatch, and other inseason reporting needs are solicited from the area management biologist. Reporting schedules are established for observers. In addition, requests for observer data collection from state of Alaska and federal Crabs in Cold Water Regions: Biology, Management, and Economics 703 agencies and national and international universities are coordinated through the ADFG research staff. The North Pacific Observer Training Center (OTC), operated by the Uni- versity of Alaska, began training observers in 1991, and since 1993 all crab observers have been trained for observer companies in a 2-week OTC course. Up to four training classes are held annually. The observer program staff also coordinates scheduling of observer training classes with OTC and observer companies, and updates OTC staff on changes in the department’s crab and scallop observer programs. The ADFG shellfish research staff manages the observer information databases. Nearly all data collected by crab observers are compiled in the Dutch Harbor office. Data are edited and electronic databases are created and archived. The final data analysis is completed by shellfish research staff in the Kodiak ADFG regional office. An annual summary of data collected by observers is published and distributed to the observers and to vessels that carried observers. Since 1993 virtually every regulation pertaining to BSAI king and Tan- ner crab fisheries or the commissioner’s permit fisheries have largely been informed by observer data. More recently, observer requirements have become important to attain the conservation, research, and management objectives of the FMP, and to fulfill required provisions of the Magnuson- Stevens Act. Also, assessing bycatch of crabs in other shellfish fisheries is an integral component to rebuilding plans for crab stocks such as Bering Sea Tanner crabs. Fishery data collected by at-sea observers provide information that could not be obtained otherwise, especially for unsurveyed or poorly sur- veyed crab stocks. Some of the biological data collected by crab observers include:

• Species catch composition from sampling the contents of randomly selected crab pots retrieved by the vessel.

• Fishing effort, harvest, and location.

• Male and female crab size measurements and shell ages.

• Chronicling reproductive cycles of crabs.

• Recording levels of incidental bycatch.

The annual number of crab observers in the program has ranged from 35 to 119. There are also five seasonal fishery biologist staff members that deploy as at-sea fishery observers in selected fisheries. The annual number of observer days has ranged from 3,672 in 1989 during the first full year of the program, to over 10,000 observer days in 1991 (Table 1). 704 Boyle and Schwenzfeier — Alaska Shellfish Observer Program

Conclusion Data collected by at-sea observers have become an integral component of the department’s BSAI shellfish management and research programs. The Shellfish Observer Program has been strengthened and improved by revisions made during the BOF process. The existing system where industry is a client of observer companies places economic pressures on observer companies that could undermine the objectivity of the observer program. Using cost recovery funds to pay direct costs of observer deployments from portions of the crab fishing vessels and deploying ADFG biologists to collect data are among the latest changes that have strengthened the state’s crab observer program. In the future we will see continued im- provements to ensure observer data quality and integrity.

Acknowledgment This is contribution PP-206 of the Alaska Department of Fish and Game, Commercial Fisheries Division, Juneau.

References Morrison, R. 1992. Eyes at sea: Alaska’s Mandatory Shellfish Observer Program. Alaska’s Wildlife 24(2):21–24. North Pacific Fishery Management Council (NPFMC). 1998. Fishery management plan for Bering Sea/Aleutian Islands king and Tanner crabs. North Pacific Fish- ery Management Council, Anchorage, Alaska. Schmidt, D., and B.A. Johnson. 1988. A comparison of catcher-processor and catcher vessel fishing performance in the 1987 Bering Sea red king crab fishery. Alaska Department of Fish and Game, Division of Commercial Fisheries, Re- gional Information Report 4K88-14, Kodiak. 19 pp. Schmidt, D., and B.A. Johnson. 1989. A comparison of catcher-processor and catcher vessel fishing performance in the 1988 Bering Sea red king crab fishery. Alaska Department of Fish and Game, Division of Commercial Fisheries, Re- gional Information Report 4K89-1, Kodiak. 13 pp. Schmidt, D., and B.A. Johnson. 1990. A comparison of catcher-processor and catcher vessel fishing performance in the 1989 Bering Sea red king crab fishery. Alaska Department of Fish and Game, Division of Commercial Fisheries, Re- gional Information Report 4K90-2, Kodiak. 8 pp. Crabs in Cold Water Regions: Biology, Management, and Economics 705 Alaska Sea Grant College Program • AK-SG-02-01, 2002

Development and Management of Crab Fisheries in Shetland, Scotland

Ian R. Napier North Atlantic Fisheries College, Port Arthur, Scalloway, Shetland, United Kingdom

Abstract Shetland is an island community forming the most northerly part of Scot- land. From the late 1950s a commercial fishery for the edible crab (Cancer pagurus) developed rapidly in Shetland, fueled by new markets and by a transfer of fishing effort from more traditional fisheries. The 1970s saw a steady decline in crab landings due to a transfer of fishing effort back to demersal fisheries. The last two decades, however, have seen a fairly steady increase in crab landings, which reached almost 600 t in 1997. This new growth resulted primarily from an increase in the number of purpose- built crab-fishing vessels. Today almost 70% of the Shetland fishing fleet targets crabs and lobsters. Most of these vessels are under 10 m in length and most are operated by one man, often on a part-time basis. In the last decade crab catches have been supplemented by the previously unexploited velvet crab (Necora puber) and shore crab (Carcinus maenas). The steady increase in crab catches and fishing effort during the 1980s and 1990s gave rise to considerable local concern about the sustainability of this fishery. In 1995, the local fishermen’s association and a number of other interested local organizations came together to seek legal powers to manage the local fisheries for crabs and other shellfish species. These powers were eventually granted by the Scottish Government in 2000, making Shetland the first community in Scotland granted powers to manage a local fishery. Initial management measures focus on controlling levels of fishing effort, and a scientific program is being established to provide managers with necessary information on the status of shellfish stocks. 706 Napier — Crab Fisheries in Shetland

Historical Background Shetland is a group of about 100 islands (15 inhabited) lying some 160 km north of the U.K. mainland and about 400 km west of Norway (Fig. 1) with a population of about 22,500 people. Administratively Shetland forms an island region (equivalent to a county) with its own local authority, the Shetland Islands Council, within Scotland, itself part of the United King- dom. Shetland (with the rest of Scotland) was ruled by the U.K. parliament in London until 1999 when many (but not all) legislative powers were transferred to the new Scottish parliament in Edinburgh. Fish catching has a long tradition in Shetland, having been practiced on a commercial basis since at least the fourteenth century, when German and northern European merchants first visited the islands to barter for dried salt fish, and on a subsistence basis for 10 times as long (probably since the islands were first settled). An important factor in the islander’s exploitation of the sea has been the long seafaring heritage handed down from the Norsemen who ruled Shetland between the ninth and fifteenth centuries and whose influence in the islands remains strong to this day. Following centuries of slow and hesitant progress, the twentieth century saw rapid advances in the islands’ fish catching industry, and today Shet- land has a modern and diverse fishing industry. The importance of fish catching to Shetland stems from the islands’ location at the heart of the rich northern European fishing grounds, but historically the islanders were probably driven to exploit the seas around them as much from necessity as from inclination. Shetland’s generally infertile soils and cool, wet, and windy climate are not favorable to agriculture, but with the addition of the sea’s resources it was possible to make a modest living. This blending of two occupations, fishing and crofting (small scale subsistence farming), created the crofter-fisherman, who dominated Shetland life until comparatively recently. At the turn of the millennium Shetland is perhaps best known to the outside world for its association with North Sea oil and in particular as the site of Europe’s largest oil terminal, at Sullom Voe, where oil brought ashore through pipelines is shipped all over the world. Despite the high profile of North Sea oil, and the undoubted economic benefits it has brought the islands, the fisheries industry continues to play a key role in the islands’ economy. In 1998 the fisheries industries had a turnover of some $260 million US, about one-third of the total value of the Shetland economy, roughly equally split between the three industry sectors of fish catching, aquaculture, and fish processing (Table 1). Fishing is important to Shetland not just because of its current contribution to the local economy but also because in these remote and resource-poor islands it is perhaps the only industry that offers a realistic potential of making a substantial long-term contribution to the local economy. Throughout the 1980s and 1990s the value of the fisheries sector grew steadily, while most other sectors of the economy either declined Crabs in Cold Water Regions: Biology, Management, and Economics 707

Figure 1. Shetland forms the most northerly part of the United Kingdom, lying roughly equidistant from Norway, Faroe, and the U.K. mainland.

Table 1. The Shetland economy in 1998.

Turnover % of total

(million $US) turnover

Fisheries industries 260.0 32.6 Fish catching 91.0 11.4 Fish processing 86.7 10.9 Aquaculture 82.3 10.3 Services 256.7 32.2 Local government 152.1 19.1 Oil industry 85.0 10.7 Agriculture 18.1 2.2 Tourism 17.6 2.2 Knitwear 7.9 0.0 Total 797.5

SIC 1999 708 Napier — Crab Fisheries in Shetland

(e.g., the oil industry) or remained at a similar level (e.g., agriculture, knitwear, and tourism). While fishing remained important in Shetland it declined in much of the rest of the United Kingdom with the result that Shetland has become one of the country’s main fishing centers. The development of crab and other shellfish fisheries in Shetland in the 1960s was a significant development for the islands and for their fishermen. It provided new opportunities for, and helped raise the average real income of, existing local fishermen and also provided additional income for a large number of crofters (small-scale farmers) and others on a part-time basis. Between 1957 and 1967 the total number of fishermen in Shetland declined slightly from about 900 to about 830 but over the same period the number of shellfish fishermen more than trebled (from under 150 to over 450), of which full-timers increased from 0 to nearly 100 (the total population of Shetland was about 19,000 in 1961) (Goodlad 1971). Today crab fishing remains an important sector of Shetland’s fishing industry. Although its contribution to the industry in terms of weight or value appears small, it accounts for a high proportion of the local fishing fleet. Crab and lobster fishing in Shetland today has many analogies with an older pattern of fishing activity which, prior to the development of shellfish fishing, had all but disappeared from the islands. The traditional Shetland fisherman was a crofter-fisherman, for whom fishing for demersal species as a part-time activity supplemented the living he could make from agriculture. This fishing activity was scattered throughout the islands. The twentieth century saw a shift away from part-time fishing as it became more of a full-time, year-round activity. Some men chose to become full-time fishermen, some focused on agriculture, while others emi- grated or joined the merchant navy. As the fishing vessels became larger and more sophisticated fishing tended to become concentrated in the relatively few communities that could offer safe harbors and the necessary facilities. The advent of shellfish fishing in the late 1950s and early 1960s created new opportunities in Shetland for part-time commercial fishing and for commercial fishing with small boats. This fishery has thus recreated a niche within Shetland’s fishing industry which had all but disappeared. Shellfish fishing today is dominated by small boats and by part-time fishermen (although today few are crofters), and is scattered throughout the islands, wherever a pier or harbor suitable for a small boat exists. Shetland’s fleet of some 250 fishing vessels ranges from shellfish fishing vessels less than five meters in length to pelagic trawlers of over 50 m. In 1998 the Shetland fleet caught about 125,000 t of fish, worth some $74 million US. White fish (demersal species such as cod, haddock, whiting, etc.) accounted for 20% of the catch (by weight), while pelagic species (herring and mackerel) accounted for 60%. Fifteen percent of the catch was industrial species (blue whiting, sandeels, etc.) and the remaining 5% was shellfish (crabs, lobsters, scallops, etc.). In addition to the catch by Crabs in Cold Water Regions: Biology, Management, and Economics 709 the local fleet, an additional 55,000 t of fish, worth some $16 million, was landed in Shetland by non-Shetland vessels in 1998. Shellfish fishing has provided a route into commercial fishing for many new, young fishermen. As demersal and pelagic fishing vessels have become larger and more sophisticated (and so more expensive) it has become increasingly difficult for new crews to raise the necessary capital to purchase even a second-hand vessel. This has been exacerbated by the increasing price of fish quota and by the increasingly tight regulation of demersal and pelagic fisheries. With relatively small vessels, and no necessity to purchase quota, the cost of entry to shellfish fisheries has remained relatively low. The ease of access to shellfish fisheries in Shetland has, however, not been without its problems. In particular it has led to these fisheries being viewed by many as “standby” occupations; occupations that can be pursued when returns from other fisheries are poor or in the absence of other employment. The last few decades have, however, seen a fundamental change in the status of Shetland’s shellfish fisheries. When they were new and stocks were unexploited their capacity to absorb new entrants and increased effort was large. Since then a relatively large permanent shellfish fishing fleet has grown up and stocks of the main shellfish species, including the edible crab, have probably been exploited to, if not beyond, their maximum sustainable levels. Given this, the capacity of shellfish fisheries to absorb new entrants or increased effort must now be all but exhausted and the view of shellfish fishing as a “standby” activity that can be entered and left at will, according to other economic factors, is no longer tenable.

Development of the Crab Fishery Although Shetlanders have exploited certain shellfish species, principally mollusks, on a subsistence basis for at least 5,000 years, and although demersal and pelagic fish species have been fished commercially for at least 500 years, the commercial exploitation of shellfish has a much more recent history, having developed largely within the last 50 years (Goodlad 1971, Nicholson 1999). The local crab fishery had its origins in the fishery for lobsters (Homarus gammarus), which expanded rapidly in Shetland in the late 1950s and early 1960s (Fig. 2). Fishermen caught edible crabs (Cancer pagurus) as bycatch with lobsters but the crabs were initially regarded as having no value and most were discarded. A major obstacle to the development of a crab fishery was the absence of local markets and the difficulty of shipping them alive to markets outside Shetland, as was (and is) common practice with lobsters. Edible crabs in close proximity invariably fight and it is difficult to tie their claws securely to prevent this (as can be done with lobsters). As a result crabs shipped alive suffered a high mortality rate in transit. Crabs also had a relatively low value compared to lobsters ($58 US per metric ton (t) compared to $876 per t in the early 1960s) and were consequently 710 Napier — Crab Fisheries in Shetland

Figure 2. Annual landings (metric tons) of edible crabs (Cancer pagurus), velvet crabs (Necora puber), and lobsters (Homarus gammarus) in Shetland, 1953-1997 (Goodlad 1971; SIC 1975-1999, SOAEFD 1979-1999, SERAD 2000).

less productive to fish, handle, and transport. Crabs were thus less attractive than lobsters at a time when the latter were abundant. The early 1960s, however, saw a number of factors come together which led to the rapid development of a targeted fishery for edible crabs. One of the most important factors was the advent of crab processing by local shellfish processing factories which cooked the crabs and dressed and froze the meat. This created a local market for crabs and thus obviated the problems associated with shipping live crabs to markets outside Shetland. Initially this led to the crabs taken as a bycatch in the lobster fishery being landed and sold, but from about 1963 crabs were fished intentionally. Lobster catches declined rapidly after 1962, apparently due to overfishing. While lobsters became scarcer it was clear that there were abundant stocks of crabs in the waters around Shetland and with local markets now established interest in exploiting them grew, and the fishery developed rapidly. By 1964, 447 t of edible crab were landed in Shetland, compared to 43 t in 1962 and just 4 t in 1961 (Fig. 2). Interest in the new crab fishery, and the lobster fishery before it, was enhanced by a depression (due to a scarcity of fish) which affected the white fish fishery in the late 1950s and early 1960s. Poor catches of white fish encouraged some fishermen to turn their attentions initially to lobster fishing, often using converted white fish fishing vessels. Others obtained new, purpose-built vessels for the new fishery. As the lobster fishery Crabs in Cold Water Regions: Biology, Management, and Economics 711 declined and the prospects for crab fishing improved effort was transferred to the new fishery. The rapid development of the crab fishery was assisted by the fact that crabs could be caught using essentially the same vessels and fishing gear as used in the lobster fishery. Financial assistance for some crab fishing boats was also provided by a trust fund established in 1964 by Christian Salvesen & Co. to benefit Shetland islanders who had worked for the company in the antarctic whaling (which was coming to an end at that time) (Nicholson 1999). Catches of crabs fluctuated but remained generally high throughout the 1960s, but the 1970s saw a period of decline. Most Shetland shellfish fisheries experienced significant declines in landings during this period, apparently as a result of increasing white fish prices which encouraged fishermen to switch from shellfish to demersal fisheries.

Crab Fishing Today Landings of edible crabs reached a low of about 100 t in 1982, but since then have increased fairly steadily (Fig. 2) and since the mid-1990s have exceeded 400 t per annum with a high of 570 t in 1997. Velvet crabs (Necora puber) have also been landed commercially since 1987. The development of this fishery mirrored in many aspects the development of the edible crab fishery in the 1960s. Fishermen previously discarded the bycatch of velvet crabs taken in the lobster fishery because no markets existed. De- velopment of market outlets since the late 1980s, however, led to recogni- tion of velvet crabs as a resource in their own right and to the development of a targeted commercial fishery for this species. Landings of velvet crabs approached 100 t in 1999 (Fig. 2). Shore crabs (Carcinus maenas) have also occasionally been fished commercially around Shetland in recent years but landings remain very small and irregular. The shore crab fishery has again been strongly influenced by the availability of market outlets. Land- ings of edible and velvet crabs in Shetland in 1999 represented about 6% and 5%, respectively, of total Scottish landings of these species. Compared to the large whitefish and pelagic sectors of Shetland’s fishing industry, crab fishing appears relatively insignificant. In 1999 crabs accounted for less than 1% of all landings by weight, although over 2% by value (Table 2). However, crab (and lobster) fishing is an important activity for the smaller vessels which make up a large proportion of Shetland’s fishing fleet. Three-quarters of the fleet is under 10 m in length and 90% of these fish for crabs and/or lobsters (Table 3). The expansion of crab fishing throughout the 1980s and 1990s has seen a proliferation in the fleet of modern, purpose-built, fast creel fishing vessels which have largely replaced the older and more traditional vessels. A high proportion of these vessels are operated on a part-time basis by persons with other jobs, either onshore or at sea, and many operate only seasonally. Most of the vessels are worked by only a single person, although the larger ones often carry an additional crewman. 712 Napier — Crab Fisheries in Shetland

Table 2. Weight and value of fish landings into Shetland by U.K. vessels (1999) and composition of Shetland fishing fleet (1998) by species group.

Weight Value No. of Landings (thousand t) (thousand $US) fishing vessels

Demersal 17,300 17,002 51 Pelagic 53,678 8,443 9 Crabs 545 626 Lobsters 10 147 Crabs and lobsters 171 combined Other shellfish 1,119 2,206 20 Total 72,652 28,424 251

SOAEFD 1999, SERAD 2000

Table 3. The size composition of the Shetland fishing fleet and of its crab and lobster fishing vessels in 1998.

Crab and lobster

Total fleet fishing vessels

Vessel No. Proportion No. Proportion of length (m) vessels of fleet vessels size class

< 5 7 3 7 100 5-7.5 84 33 78 93 7.5-10 95 38 83 87 10-20 32 13 3 9 > 20 33 13 0 0 Total 251 171 68

SOAEFD 1999 Crabs in Cold Water Regions: Biology, Management, and Economics 713

Shetland’s crab fishery remains closely associated with the lobster fishery from which it developed; most vessels that fish for crabs also fish for lobsters and vessels targeting lobsters also take crabs as bycatch. It is thus impractical, and to a large extent meaningless, to attempt to draw a clear division between the crab and lobster fisheries. Both crabs and lobsters are fished using creels (traps), generally set on “leaders,” or lines that join the individual creels. The creels are rectangular and semicircular in cross-section and generally range from about 0.6 to 0.9 m in length by about 0.5 m in width. Traditionally they were home-made, with a wooden base and wooden, bamboo, or wire hoops covered in netting, but most are now commercially manufactured with plastic coated, welded-steel frames, again covered in netting. Crabs and lobsters gain entry via funnel shaped entrances in the sides. Crab creels tend to be somewhat larger than lobster creels, and often incorporate an inner chamber, or “parlor,” to improve retention rates. The creels are hauled mechanically on all but the very smallest fishing vessels. Lobsters and velvet crabs are generally fished close to shore on rocky or stony bottoms in depths down to about 50 m. Edible crabs are generally fished further offshore on soft (sandy) bottoms in depths down to 100 m or more. The number of creels fished and the number of creels per leader depends very much on the size of the vessel and the area being fished. Most of the regularly active vessels work between 100 and 200 creels, although the larger vessels, particularly those being operated on a full- time basis, may work up to 400 or 500 creels, with a few boats working as many as 1,000 or more. Smaller vessels may work as few as a dozen or so. The creels are generally hauled after 24 hours, although part-time fishermen or those working large numbers of creels may leave them in the water for longer. The most commonly used bait is white fish offal, usually obtained from fish processing factories. Shortage of bait, due to a downturn in the white fish fishery and processing sector has been a problem in recent years and some fishermen experiment with alternatives such as pelagic species or obtain undersized fish directly from white fish fishing vessels. In recent years there have been two main markets for crabs landed in Shetland: local processing and live transport to markets on continental Europe. Live transport has been by “vivier” trucks, which are vehicles equipped to transport live shellfish in seawater tanks. A number of such vehicles operated in Shetland (and other parts of Scotland) in the 1990s, purchasing shellfish directly from fishermen for transport to markets in continental Europe, mainly Spain. This played a key role in developing the velvet crab fishery by providing a market outlet, and it has also provided the only outlet for shore crabs landed in Shetland. Only one company in Shetland now processes crabs, including virtually all the edible crabs landed in Shetland and a variable proportion of the velvet crabs. About 50% of the edible crab is canned (cooked) for the U.K. market, with most of the remainder being produced in a variety of fresh or frozen formats including whole cooked crabs and cooked claws, 714 Napier — Crab Fisheries in Shetland and dressed crab meat. About 30% of the fresh and frozen products are sold in the U.K. market while the remainder is exported, mainly to France and Spain. Velvet crabs are frozen whole, either raw or cooked, almost en- tirely for export to Spain.

Crab Fishery Management Within the United Kingdom, crab (and other shellfish) fisheries have been largely unregulated, particularly when compared to the demersal and pelagic sectors. In particular there are no limits on fishing effort or on catches, although there are minimum landing sizes of 140 mm carapace width (CW) for edible crabs (in the northern North Sea, including Shetland) and 65 mm CW for velvet crabs. The absence of limits allowed crab fishing to expand rapidly during the 1980s and 1990s with landings in Shetland increasing more than five-fold between 1982 and 1997. There was considerable concern that if this trend were to continue, the future viability of the crab fisheries, and thus of a large proportion of the local fishing fleet, could be threatened. In the absence of national controls, the Shetland Fishermen’s Associa- tion (the representative body of Shetland’s fishermen) sought a legal mechanism by which fishing for crabs and other shellfish species might be managed locally. The only possible mechanism they were able to identify was to obtain a Regulated Fishery Order, or “regulating order.” Regulating orders can be granted within the United Kingdom by the government and confer legal rights to manage shellfish fisheries to a local organization. Regulating orders apply to the fisheries for specified shellfish species within a specified area and are intended to provide for the establishment, improvement, maintenance and/or regulation of shellfish fisheries. Al- though regulating orders were available since the 1960s the range of species which could be covered was until fairly recently very limited and only a few orders were granted, none of them in Scotland. In 1995 a group of interested parties agreed that they would seek to obtain a regulating order to cover local Shetland shellfish fisheries. To formally apply for and implement the order this group established a new organization, the Shetland Shellfish Management Organization (SSMO). The SSMO was formed as a partnership of local organizations; Shetland Fishermen’s Association, Shetland Islands Council, Shetland Association of Community Councils, Shetland Fish Processor’s Association, North At- lantic Fisheries College, and Scottish Natural Heritage. The primary aim of the organization is to manage Shetland’s shellfish fisheries to ensure their long-term sustainability. After 5 years of negotiations with the government, and a public inquiry, the order—Shetland Islands Regulated Fishery Order 1999 (Scottish Statu- tory Instruments 1999)—was finally granted, coming into force in January 2000. The order covers the waters around Shetland between the low water mark and the 6-mile limit, an area of about 6,000 km2, and applies to all Crabs in Cold Water Regions: Biology, Management, and Economics 715 shellfish species that are, or might be, fished commercially (oysters, mussels, cockles, clams, lobsters, scallops, queen scallops, crabs, whelks and razorshells). Under the order the SSMO implemented a local shellfish fisheries management plan, centered around a local licensing scheme aimed at controlling fishing effort. Certain restrictions on fishing gear and vessel sizes have also been introduced, along with minimum landing sizes and closed seasons for some species. The SSMO drew up its management plan, and continues to update it, in close consultation with the local fishing industry. It has also carried out an extensive consultation exercise providing an opportunity for all shellfish fishermen to express their views on the status of shellfish stocks and on what management measures they think would be beneficial. These close links with the industry are essential, as in the absence of strong powers of enforcement the SSMO has to manage largely by consensus. Within the first year of operation some 150 fishing vessels applied for and were granted local shellfish fishing licenses. The management plan is backed by a program of fisheries data collection and research being developed by the North Atlantic Fisheries College. This program is intended to provide the SSMO with management advice and may result in additional regulations or controls being introduced in the future. The college and SSMO are jointly developing a lobster stock enhancement program to help local lobster stocks recover from overfishing in the 1960s. The Shetland Islands Regulated Fishery Order 1999 was the first of its kind to be granted in Scotland. It represented the first time that the government granted a Scottish community local control of a local fishery. Other areas of Scotland follow Shetland’s experiences with interest and a number are developing plans for regulating orders for their own areas. The granting of the Shetland regulating order was a highly significant development, not just for Shetland but also in Scotland. Its significance lies not just in the fact that it enables controls to be introduced to a virtually unregulated sector of the fisheries industry but, perhaps more impor- tantly, in that it delegates control of local fisheries away from central government to a local community. Shetland’s shellfish fishermen have on the whole responded very positively to the order, and to the implementation of local regulation of shellfish fisheries. Most recognize the need for better management of these fisheries and are pleased to have a local organization which they feel can address their concerns and which can intro- duce controls that they feel are necessary. Regulating orders are, however, not an ideal mechanism for local control of fisheries. In particular, any measures implemented under the order remain subject to government approval. To date the mechanisms for ap- proving measures (and the order itself) have proven to be slow and somewhat cumbersome. In addition there has been reluctance on the part of the government to consider measures which do not enjoy more or less unanimous support of all fishermen. While this ensures a general consensus 716 Napier — Crab Fisheries in Shetland in the management of the fisheries it could make the implementation of stricter regulations difficult, should they prove necessary. Finally, it remains to be seen how shellfish regulating orders will fare without the government support for research, assessment, and enforcement which is provided for demersal and pelagic fisheries. One of the underlying problems with regulating orders is that the original enabling legislation was drafted in the 1960s when shellfish fishing in the United Kingdom was relatively undeveloped. The emphasis of the legislation is, therefore, largely on the development of shellfish fisheries. Today most shellfish fisheries are fully developed, if not over-developed, and the primary requirement in many cases is to reduce fishing effort. It may prove difficult to achieve this using regulating orders, particularly given the current government reluctance to approve management measures which are not unanimously supported by fishermen. These problems aside, regulating orders remain a significant development in Scotland, although the significance may lie more in the principle they establish (of delegation of fisheries management) than in the detail of how they are used. The SSMO and the Shetland shellfish regulating order face many challenges, but if successful they will help ensure the long-term sustainability of Shetland’s shellfish fisheries. In turn, this will help maintain an important sector of the local fishing industry, providing continued opportunities for small-boat, full- and part-time fishermen. As the first to be granted, the Shetland order is certain to remain the focus of much attention.

References Goodlad, C.A. 1971. Shetland fishing saga. The Shetland Times Ltd., Shetland. 343 pp. Nicholson, J.R. 1999. Shetland fishermen. The Shetland Times Ltd., Shetland. 157 pp. Scottish Statutory Instruments. 1999. The Shetland Islands Regulated Fishery (Scot- land) Order 1999. Scottish Statutory Instruments 1999 No. 194. Edinburgh, HMSO. (Available from [August 2001]). SERAD (Scottish Executive Rural Affairs Department). 2000. Scottish sea fisheries statistics 1999. The Scottish Executive, Edinburgh. SIC (Shetland Islands Council). 1975-1999. Annual Publication. Shetland in statistics. Development Department, Shetland Islands Council, Shetland. SOAEFD (Scottish Office Agriculture, Environment and Fisheries Department). 1979- 1999. Annual publication. Scottish sea fisheries statistics. The Scottish Office, Edinburgh. Crabs in Cold Water Regions: Biology, Management, and Economics 717 Alaska Sea Grant College Program • AK-SG-02-01, 2002

Mortality of Chionoecetes Crabs Incidentally Caught in Alaska’s Weathervane Scallop Fishery

Gregg E. Rosenkranz Alaska Department of Fish and Game, Division of Commercial Fisheries, Kodiak, Alaska

Abstract Catcher-processor vessels fishing for weathervane scallops (Patinopecten caurinus) off Alaska have been required to carry onboard observers on all trips since 1993. Besides collecting biological data on scallops, these observers examined over 100,000 Tanner crabs (Chionoecetes bairdi) and snow crabs (C. opilio) incidentally caught by dredges in the Bering Sea scallop fishery and classified each as dead or alive. I used graphical methods and generalized linear modeling (GLM) to explore relationships between mortality rates (proportion classified dead) and variables such as species, sex, shell condition, and injuries. Mortality rates were high for crabs with injury to the carapace and there was a strong positive relationship between mortality rate and number of new injuries. Mortality rates were consistently higher for snow crabs than for Tanner crabs, and mortality rates tended to decline with shell size for new-shell crabs of both species. Due in part to large sample sizes, all variables analyzed except weight of the scallop catch were significant in the GLM. Between-year and between-vessel differences in mortality rates were also important to GLM fit. Overall, 24% of the crabs inspected by observers were recorded as dead.

Introduction Alaska’s offshore weathervane scallop (Patinopecten caurinus) fishery is prosecuted by a small fleet of catcher-processor vessels that harvests from beds located along the northern Gulf of Alaska and in the eastern Bering Sea. Fishing is with New Bedford–style dredges and typically occurs in depths of 50-140 m. Seasons begin on July 1 each year and continue until either area-specific harvest ranges or bycatch caps are met. Statewide landings 718 Rosenkranz — Mortality of Chionoecetes Crabs of shucked meats average about 400,000 kg per year, yielding an ex-vessel value of approximately US$5 million. The statewide scallop observer program was implemented by the Alaska Board of Fisheries in 1993 to collect biological data on scallops and to monitor bycatch of other commercially important species (Barnhart 2000). Beginning in summer of 1993, all vessels harvesting scallops in Alaskan waters outside Cook Inlet have been required to carry fishery observers on all trips (Barnhart 2000). In addition to collecting biological data on scallops and identifying and weighing bycatch, the observers made detailed examinations of large numbers of Tanner crabs (Chionoecetes bairdi) and snow crabs (C. opilio) incidentally caught in the fishery. These observer-collected data provide a rich source of information on mortality of crabs incidentally caught by mobile fishing gear. This study reports on mortality of Tanner and snow crabs incidentally caught in Alaska’s scallop fishery. Relationships between mortality rates and variables such as crab size and shell condition are explored. I focus on data from the eastern Bering Sea because it is an important fishing area for Tanner and snow crabs as well as scallops; the Bering Sea scallop fishery often closes when crab bycatch caps are attained. Graphical examination of the data and generalized linear modeling were the primary tools used in the analysis.

Data and Methods Observers were deployed on all trips of all scallop catcher-processors and instructed to randomly sample one dredge from each of six hauls daily for bycatch (Barnhart 1998). During 1993-1994, all Tanner and snow crabs from each sampled dredge were examined; from 1996 to 1999, a maximum of 20 randomly selected individuals of each species from each sampled tow were examined. The scallop fishery was not opened in 1995 due to problems related to inconsistent state and federal regulations. Data collected included species, sex, shell condition, carapace width (CW), number of new injuries (new injuries), presence or absence of injury to the carapace (carapace injury), and mortality. Crushed and moribund crabs were classified as dead, and crabs that appeared to have a good chance of survival were classified as alive. Shell condition was either new, generally 1 year or less past the most recent molt and characterized by white ven- tral surfaces and relatively few scratches or abrasions on the carapace and appendages, or old (Barnhart 1998). New injuries were damage of obvi- ously recent origin distinguishable from old injuries such as autotomized or regenerated limbs. For the analysis, all C. bairdi ¥ C. opilio hybrids were considered snow crabs. Auxiliary haul-specific data available for the bycatch-sampled tows included fishing date, vessel, and estimated round weight catch of scallops (scallop catch). The observer data were first summarized for sample sizes and mortality by year, species, sex, shell condition, carapace width, carapace injury, and new injuries. Plots were constructed to compare mortality rates Crabs in Cold Water Regions: Biology, Management, and Economics 719

(proportion classified dead) for different combinations of the categorical variables. Combination histogram–line plots were used to show sample sizes and mortality rates by carapace width, new injuries, and scallop catch. A generalized linear model (GLM; McCullagh and Nelder 1989) with logit link was then fit to the data; this essentially modeled the log odds of mortality as a linear function of categorical (year, vessel, species, sex, shell condition, and carapace injury) and continuous (carapace width, new injuries, and scallop catch) variables. A stepwise fitting procedure using Akaike’s information criterion (AIC; e.g., Venables and Ripley 1994) was employed for model selection. First, a base model with all variables but no interaction terms was fit. With this model, an estimate of the dispersion parameter f was obtained from the generalized Pearson statistic (Christensen 1997, pp. 307-309). For testing further models, I used dispersion-adjusted AIC, defined as

ˆ AIC =+ Dp2 f , where D is the model deviance, and p is the number of estimated GLM parameters. I refit the model after dropping the original variables one at a time, and if the simpler model produced equal or smaller AIC, the variable was eliminated from further consideration. Interaction terms were then added one at a time, and reduction in AIC was again used as the metric for inclusion. I checked for interactions between each continuous variable and each categorical variable included in the base model. A goodness-of- fit test suggested by Christensen (1997, p. 129) was used for model check- ing. This test divides the [0, 1] interval into 10 equal-width subintervals based on predicted probabilities, then compares the observed number of mortalities for observations in the subinterval with percentiles of a binomial distribution, with n equal to the number of observations in the subinterval and p equal to the average of predicted probabilities for observations in the subinterval.

Results Graphical Analysis and Summary Statistics Annual estimates of crab bycatch in the Bering Sea scallop fishery during 1993-1999 ranged from 17,000 to 291,000 Tanner crabs and from 15,000 to 233,000 snow crabs (Fig. 1; Barnhart and Rosenkranz 2000). Of these, 60,350 Tanner crabs and 40,818 snow crabs were examined by scallop observers. For purposes of comparison, eastern Bering Sea snow crab harvests averaged 118.2 million crabs per year from 1993 to 1999 and Tan- ner crab harvests averaged 3.0 million crabs per year from 1993 to 1997, when the fishery was closed due to low abundance (ADFG 2000). Sample sizes of incidentally caught crabs varied considerably over time by species, sex, and shell condition (Fig. 2). In 1993 and 1994, when 720 Rosenkranz — Mortality of Chionoecetes Crabs

Figure 1. Estimates of crab bycatch in the Bering Sea scallop fishery since inception of the scallop observer program.

Tanner crabs were relatively abundant on the scallop fishing grounds and all crabs in each sampled dredge were examined, data on over 25,000 Tanner crabs were collected by the observers. Sample sizes for Tanner crabs decreased to around 1,700 per year from 1996 to 1999, while snow crab sample sizes peaked in 1996 and 1997, when approximately 11,000 were examined. Old-shell female and new-shell male Tanner crabs dominated the bycatch during 1993-1994, and male snow crabs were more common throughout; new-shell females composed a minor component of the snow crab bycatch, likely due to spatial distribution patterns that vary with sex and age (Zheng et al. 2001). Mortality rates of incidentally caught crabs examined by scallop observers also varied substantially over time. Relatively low values were observed for both Tanner and snow crabs in 1993 and 1996, with higher mortality rates in 1997 and 1998 (Fig. 3). Correlation between mortality rates by species was surprisingly high (r2 = 0.99), with rates for snow crabs slightly exceeding those for Tanner crabs in each year. Trends in mortality rates by sex and shell condition were not consistent. For example, female new-shell Tanner crabs experienced the highest mortality rates in 1993-1994 and the lowest rates in 1996-1999. For male snow crabs, mortality rates were similar for old- and new-shell animals in 1994, 1996, and 1998, but the old-shell mortality rate exceeded the new-shell rate by factors of 2.1 in 1993 and 3.8 in 1999. Bycatch mortality rates might also be influenced by crab size, which is intrinsically related to sex and shell condition in Chionoecetes crabs due to terminal molting (e.g., Somerton 1981, Saint-Marie et al. 1995). Plots depicting 1994 Tanner crab mortality rates by carapace width (Fig. 4) suggest declining mortality with size for small new-shelled crabs, relatively constant rates for larger new- and old-shell males, and modest increases in mortality rates with carapace width for old-shell females. Tanner crab Crabs in Cold Water Regions: Biology, Management, and Economics 721

Figure 2. Sample sizes (numbers examined by observers) for Tanner and snow crabs incidentally caught in the Bering Sea scallop fishery during 1993-1999.

data from 1993 (not shown) exhibited similar trends. Snow crab mortality rates for 1997 (Fig. 5) also tended to decrease with size, but in contrast to Tanner crabs, the trend was apparent for old-shell females as well as new- shelled crabs. However, this trend was not repeated for old-shell females in 1998, the only other year with a reasonably large sample size. From one to eight vessels participated in the Bering Sea scallop fishery each year since the inception of the observer program. Mortality rates of crabs incidentally caught by these vessels ranged from 0.02 to 0.72. Examination of mortality rates for the three vessels with the most fishing time in the area since 1993 (Fig. 6) revealed no distinct patterns of variability. For example, the snow crab mortality rate for “Vessel A” increased from 0.05 in 1993 to 0.63 in 1999, while the rate for “Vessel C” decreased from 0.30 in 1993 to 0.07 in 1999. No single vessel consistently produced 722 Rosenkranz — Mortality of Chionoecetes Crabs

Figure 3. Mortality rates (proportion classified as dead) for Tan- ner crabs (see Fig. 2 legend); snow crabs (see Fig. 2 legend); and Tanner (solid line) and snow (dashed line) crabs incidentally caught in the Bering Sea scallop fishery during 1993-1999. Crabs in Cold Water Regions: Biology, Management, and Economics 723

Figure 4. Carapace width distributions and mortality rates for new-shell male, old-shell male, new-shell female, and old-shell female Tanner crabs incidentally caught and sampled in the 1994 Bering Sea scallop fishery. 724 Rosenkranz — Mortality of Chionoecetes Crabs

Figure 5. Carapace width distributions and mortality rates for new-shell male, old-shell male, and old-shell female snow crabs incidentally caught and sampled in the 1997 Bering Sea scallop fishery. New-shell females are not depicted due to small sample size. Crabs in Cold Water Regions: Biology, Management, and Economics 725

Figure 6. Mortality rates of incidentally caught Tanner and snow crabs for the three vessels with the most fishing effort in the Ber- ing Sea scallop fishery during 1993-1999. 726 Rosenkranz — Mortality of Chionoecetes Crabs the highest or lowest mortality rate for either species. Note that different observers were deployed on the vessels each year. Injuries clearly affect the chances of survival for crabs incidentally captured in scallop dredges. Within the scallop observer data set, injuries to the carapace were relatively rare but lethal. About 5% of incidentally caught crabs sustained carapace injury, and of these, about 85% died (Table 1). Relationships between new injuries and mortality were also relatively consistent, with more injuries leading to higher mortality rates (Fig. 7). The majority of incidentally caught crabs received no injuries; however, 8% of uninjured Tanner crabs and 24% of uninjured snow crabs were classified as dead by the observers. Plots of mortality rate versus scallop catch revealed no relationships whatsoever.

GLM Fitting the GLM quantified results of the graphical analysis: probability of mortality for crabs incidentally caught in the Bering Sea scallop fishery varied significantly by species, sex, shell condition, carapace width, new injuries, carapace injury, year, and vessel. Scallop catch was the only variable without an effect on mortality. The model selected by the stepwise procedure (Table 2) contained 54 parameters, including 33 interaction terms, and produced residual deviance of 28,884 on 24,653 degrees of freedom. Values of t-statistics for the estimated GLM coefficients indicated that the effects of carapace injury and new injuries were extremely significant, species and shell condition were moderately significant, and carapace width and sex were less significant. Inclusion of interaction terms involving carapace width and sex reduced absolute values of t-statistics for carapace width from 4.6 to 2.4 and for sex from 14.0 to 1.5. The effects of year and vessel were also important to the model fit. Interactions were significant between the two continuous variables (new injuries and carapace width) and a number of the categorical variables. For each continuous variable, the GLM fits a linear predictor for each level of each categorical variable; interaction terms allow these lines to have different slopes. Hence, slopes of predicted linear relationships between number of new injuries and log odds of survival differed for each level of shell condition, species, carapace injury, vessel, and year, but not for sex. For carapace width, slopes differed for each level of sex, species, vessel, and year, but not for shell condition or carapace injury. As an example, the effect of interaction between new injuries and vessel for new- shell male Tanner crabs in 1993 is shown in Fig. 8. Slopes of the lines that predict log odds of mortality are quite different for the two vessels, leading to contrasting shapes of the logistic curves that predict probability of mortality. Crabs in Cold Water Regions: Biology, Management, and Economics 727

Table 1. Sample sizes and mortality rates of observer-sampled Tanner and snow crabs that received injury to the carapace when incidentally caught in the Bering Sea scallop fishery.

Carapace injury Carapace injury and mortality

Proportion

N N Proportion N of injured

New-shell Tanner crabs 24,798 1,525 0.061 1,340 0.879 Old-shell Tanner crabs 35,552 1,059 0.030 864 0.816 New-shell Snow crabs 22,446 1,136 0.051 961 0.846 Old-shell Snow crabs 18,372 880 0.048 748 0.850

Figure 7. Sample sizes and mortality rates by number of new injuries for observer-sampled Tanner and snow crabs incidentally caught in the Bering Sea scallop fishery during 1993-1999. 728 Rosenkranz — Mortality of Chionoecetes Crabs

-value

statistics for the GLM.

alue S.E.

-value Parameter V

Absolute Absolute t

Presence or absence of carapace injur

Carapace width.

alue S.E.

0.885 0.034 26.362 NewInj:VesselF 0.151 0.042 3.604 0.004 0.002 2.437 NewInj:VesselG 0.389 0.076 5.150

3.668 0.057 64.825 NewInj:Year94 –0.384 0.029 13.149

–0.059 0.023 2.568 CW:Year94 –0.005 0.001 4.719

Parameter estimates, standard errors, and absolute values of

sselD 0.785 0.208 3.767 NewInj:Year98 –0.671 0.041 16.528

able 2.

esselB –1.033 0.213 4.849 NewInj:Year96 0.492 0.057 8.570 esselC –2.103 0.342 6.152 NewInj:Year97 –0.662 0.039 16.809

esselE –0.727 0.272 2.668 NewInj:Year99 –0.718 0.039 18.285 esselF 0.123 0.206 0.600 CW:Sex 0.003 0.001 2.659 esselG –2.180 0.407 5.357 CW:Species –0.008 0.001 7.626 esselH –1.347 0.259 5.201 CW:VesselB 0.003 0.002 1.438 esselI –2.141 0.290 7.387 CW:VesselC 0.007 0.003 2.300 esselJ –4.500 0.247 18.238 CW:VesselD –0.007 0.002 3.970

ear94 1.841 0.119 15.498 CW:VesselE –0.003 0.002 1.349 ear96 2.974 0.249 11.947 CW:VesselF 0.002 0.002 0.997 ear97 3.015 0.157 19.213 CW:VesselG –0.004 0.004 0.864 ear98 1.806 0.183 9.871 CW:VesselH 0.001 0.002 0.511 ear99 3.094 0.192 16.086 CW:VesselI 0.007 0.002 2.840

Number of new injuries.

Parameter

Intercept –3.328 0.186 17.858 NewInj:VesselE 0.109 0.052 2.114 NewInj

CW Sex 0.139 0.094 1.479 NewInj:VesselH 0.382 0.056 6.787 Shell –0.159 0.027 5.816 NewInj:VesselI 1.022 0.069 14.753 Species 1.328 0.105 12.684 NewInj:VesselJ 0.541 0.046 11.863 CarInj V V Ve V V V V V V Y Y Y Y Y NewInj:Shell –0.083 0.019 4.369 CW:VesselJ 0.024 0.002 11.003 NewInj:Species NewInj:CarInj –0.337 0.033 10.099 CW:Year96 –0.016 0.002 6.313 NewInj:VesB 0.407 0.042 9.585 CW:Year97 –0.009 0.001 5.861 NewInj:VesCNewInj:VesD 0.344 0.231 0.068 0.044 5.026 5.305 CW:Year98 CW:Year99 0.001 –0.013 0.002 0.002 0.718 6.704 a Crabs in Cold Water Regions: Biology, Management, and Economics 729

Figure 8. Linear (top) and logistic curve (bottom) GLM predictions of mortality as a function of number of new injuries for male new-shell Tanner crabs incidentally caught in the 1993 Ber- ing Sea scallop fishery. Predictions assume 125 mm CW and no carapace injury. 730 Rosenkranz — Mortality of Chionoecetes Crabs

The goodness-of-fit test confirmed the presence of over-dispersion or extra-binomial variability within the data set. After dividing all observations into intervals based on probabilities of mortality predicted by the GLM, the observed number of mortalities fell outside the central 95% of the binomial density for 7 of the 10 intervals. Estimates of the dispersion parameter were 2.65 for the base model and 1.90 for the final model.

Discussion The main conclusion of this analysis is that a large proportion of Chionoecetes crabs incidentally caught in scallop dredges survive capture, at least initially. Of more than 100,000 crabs examined by scallop observers in the Bering Sea from 1993 to 1999, 24% were classified as dead. This is considerably lower than the rate found by Stevens (1990), who studied survival of Bering Sea crabs incidentally caught in groundfish trawls. Of 16,498 Tanner crabs captured during his 1-month study, 98% were classified as dead or moribund upon first examination; however, only 35% of a subsample of moribund crabs held in live tanks for 48 hours died, yielding an overall estimated mortality rate of 78%. Tow length and catch are much different in the groundfish trawl and scallop dredge fisheries. In Stevens’ (1990) study, average catch per tow was 20 t (primarily yellowfin sole, Limanda aspera; rock sole, Lepidopsetta bilineata; and Pacific cod, Gadus macrocephalus), and tows ranged from 1 to 6.4 hours. For the scallop fishery, catches average about 500 kg of scallops (round weight), and tow duration averages about 1 hour. These factors undoubtedly account for much of the difference in mortality rates. Unobserved mortality may increase the effect of the scallop fishery on the crab population in at least two ways. First, incidentally caught crabs returned to the ocean may die due to injuries or face higher susceptibility to predation than uninjured animals. Second, uncaptured crabs may be injured by dredges. Caddy (1973) noted that predators were attracted by the passage of scallop dredges, apparently to feed on animals damaged but not captured by the gear. Presumably, these predators also would feed opportunistically on injured or sluggish crabs returned to the sea from scallop vessels. In addition, the effects of dredging on Tanner and snow crab habitat are unknown. Scallop dredging occurs in a relatively small portion of Bering Sea crab habitat (Somerton 1981, Barnhart and Rosenkranz 2000, Zheng et al. 2001), and crab mortality due to scallop dredging is believed to be a minor component of total mortality, which is dominated by natural- and fishing-mortality rates (Witherell and Harrington 1996). Carapace injury and new injuries produced the most consistent, in- tuitive relationships with mortality rates. Carapace injury was the best predictor of mortality, and increases in the number of new injuries led to relatively linear increases in mortality rates. There was also a fairly consistent trend of decreasing mortality with size for small crabs, but relationships between mortality rates and most of the variables changed over Crabs in Cold Water Regions: Biology, Management, and Economics 731 time. Changes in sample sizes and the size-age structure of the crab populations undoubtedly contributed to this variability. Differences in classification of dead and live crabs by individual observers may also be a factor. Lack of a relationship between crab mortality rate and scallop catch, and the relatively high mortality rates for uninjured crabs, were two somewhat surprising results from the analysis. Observers have reported that crabs are crushed on deck when the catch is dumped from the dredge. This would seem more likely to occur when scallop catches are large, but I found no evidence of increasing mortality with increasing scallop catch in the data. Several explanations may be relevant to mortality of crabs recorded as uninjured. Damage to mouthparts and minuscule cracks in the carapace are two potentially fatal injuries that are not likely to be reported by observers. Sediment suspended by the dredge may clog the gills (A.J. Paul, University of Alaska Fairbanks, Institute of Marine Science, pers. comm.), producing another type of injury that would not be recorded. Finally, differences in classification of dead and live crabs by individual observers may again be a factor. Over-dispersion is a relatively common phenomenon for binary-data GLMs (McCullagh and Nelder 1989). Results of the goodness-of-fit test and estimates of dispersion parameters indicating its presence were not unex- pected; the graphical analysis showed that relationships between explana- tory variables and mortality rates were highly variable and not always consistent. The GLM with binomial error could not account for all the variability within the large scallop observer data set. I assumed that mean values such as the GLM parameters were not affected, but that variance was underestimated. After correcting for this problem by using dispersion-adjusted AIC for model selection, model inference was likely not affected by over-dispersion.

Acknowledgments Special thanks are due to J.P. Barnhart of ADFG, coordinator of the weathervane scallop observer program, and to dozens of observers who collected the data. I also thank Doug Pengilly and other reviewers for their comments. This is contribution PP-208 of the Alaska Department of Fish and Game, Commercial Fisheries Division.

References Alaska Department of Fish and Game (ADFG). 2000. Annual management report for the shellfish fisheries of the westward region, 1999. Alaska Department of Fish and Game, Division of Commercial Fisheries, Kodiak, Regional Informa- tion Report 4K00-55. 345 pp. Barnhart, J.P. 1998. Weathervane scallop observers manual. Alaska Department of Fish and Game, Division of Commercial Fisheries, Regional Information Re- port 4K98-32, Kodiak. 44 pp. 732 Rosenkranz — Mortality of Chionoecetes Crabs

Barnhart, J.P. 2000. Weathervane scallop fishery in the westward region, 1967-1999: Report to the Alaska Board of Fisheries. Alaska Department of Fish and Game, Division of Commercial Fisheries, Regional Information Report 4K00-12, Kodi- ak. 40 pp. Barnhart, J.P. and G.E. Rosenkranz. 2000. Summary and analysis of onboard observer-collected data from the 1998/99 statewide commercial weathervane scallop fishery. Alaska Department of Fish and Game, Division of Commercial Fisheries, Regional Information Report 4K00-812, Kodiak. 86 pp. Caddy, J.F. 1973. Underwater observations on tracks of dredges and trawls and some effects of dredging on a scallop ground. J. Fish. Res. Board Can. 30:173- 180. Christensen, R. 1997. Log-linear models and logistic regression. Springer, New York. 483 pp. McCullagh, P., and J.A. Nelder. 1989. Generalized linear models. Chapman and Hall/ CRC, New York. 511 pp. Saint-Marie, B., S. Raymond, and J. Brêthes. 1995. Growth and maturation of the benthic stages of male snow crab, Chionoecetes opilio (Brachyura: Majidae). Can. J. Fish. Aquat. Sci. 52:903-924. Somerton, D.A. 1981. Regional variation in the size and maturity of two species of Tanner crab (Chionoecetes bairdi and C. opilio) in the eastern Bering Sea, and its use in defining management subareas. Can. J. Fish. Aquat. Sci. 38:63-174. Stevens, B.G. 1990. Survival of king and Tanner crabs captured by commercial sole trawls. Fish. Bull., U.S. 88:731-734. Venables, W.N., and B.D. Ripley. 1994. Modern applied statistics with S-Plus. Springer- Verlag, New York. 462 pp. Witherell, D., and G. Harrington. 1996. Evaluation of alternative management measures to reduce the impacts of trawling and dredging on Bering Sea crab stocks. In: High latitude crabs: Biology, management, and economics. University of. Alaska Sea Grant, AK-SG-96-02, Fairbanks, pp. 42-58. Zheng, J., G.H. Kruse, and D.R. Ackley. 2001. Spatial distribution and recruitment patterns of snow crabs in the eastern Bering Sea. In: G.H. Kruse, N. Bez, A. Booth, M.W. Dorn, S. Hills, R.N. Lipcius, D. Pelletier, C. Roy, S.J. Smith, and D. Witherell (eds.), Spatial processes and management of marine populations. University of Alaska Sea Grant, AK-SG-01-02, Fairbanks, pp. 233-255. Crabs in Cold Water Regions: Biology, Management, and Economics 733 Alaska Sea Grant College Program • AK-SG-02-01, 2002

Occurrence of Northern Stone Crab (Lithodes maja) at Southeast Greenland

Astrid K. Woll Møre Research, Ålesund, Norway

AnnDorte Burmeister Greenland Institute of Natural Resources, Nuuk, Greenland

Abstract A joint Norwegian-Greenlandic trial pot fishery for northern stone crab (Lithodes maja) was conducted in the summer of 1995 and 1996. The objectives were to evaluate the occurrence of L. maja according to area, depth, and temperature. Catch rates, sex, and size distribution and fecundity of females were investigated to gain a better understanding of the resource potential. In 1995 the fishery was conducted on the continental shelf (62º- 63ºN) and in Ammasalik Fjord (65ºN). A total of 763 crabs were caught. Catch rates for crabs of commercial size (more than 0.6 kg) were estimated

(CPUEc) and were 0.23 kg per pot haul on average for all localities. Highest

CPUEc was found on the continental slope on steep and rocky bottom and temperature about 4-5ºC (62ºN). In 1996 a trial fishery was conducted from the slope (62ºN) and westward to the Tingmiarmiut Fjord. A total of 1,134 crabs were caught and overall CPUEc was 0.81 kg per pot haul. Crabs more than 0.6 kg were processed and frozen onboard and used for market testing in restaurants in Norway, which concluded that the meat was of excellent quality, but a better price would be obtained for a live or fresh product. Males were bigger than females and 98.6% of the crabs more than 0.6 kg were males. Females caught were in the size range of 45-125 mm carapace length (CL) and a total of 51.9% of these had egg clutches. Mean egg diameter was 2.16 mm and fecundity was positively correlated to CL.

Introduction Several species of crabs are known to occur in the waters surrounding Greenland. Only the snow crab (Chionoecetes opilio) is commercially exploited. 734 Woll and Burmeister — Northern Stone Crab

Two other species may also be of commercial interest, the northern stone crab (Lithodes maja) and the porcupine crab (Neolithodes grimaldi) (Squires 1990). These species have occasionally been taken as bycatch in trawls on the banks off the west coast of Greenland, L. maja at depths between 50 and 800 m and N. grimaldi between 500 and 2,000 m (Andersen 1993). L. maja is distributed in the North Atlantic, from Labrador to New Jersey, along the coast of west and east Greenland, along the coasts of Iceland and Spitzbergen, and along the Norwegian coast to the British Isles and the Netherlands. Until now there has been no direct target fishery for the crab. In Canada a substantial bycatch of L. maja was reported by gillnet fishermen along the coast of Newfoundland, and in 1992 the Department of Fisheries of Newfoundland and Labrador carried out a trial fishery (Dooley and Johnsen 1993). Trial fisheries were also conducted in 1994/1995 (DFOC 1995) and 1996 (CAFID 1996) in approximately the same areas. Resources of commercial interest were found, but processing and marketing were difficult. In Norway L. maja has been reported as a bycatch in several areas. In northern Norway a trial fishery was conducted in 1992 and 1993 (Hufthammer 1996). The crabs were processed as clusters which were single frozen and glazed (Brataas 1993). Market testing was carried out in Germany, concluding that the size of the crab should be more than 0.6 kg (live weight) (Bratteng 1994). Joint Norwegian-Greenlandic trial fisheries started in 1994 as part of a mapping of Greenlandic resources at the southeast coast. In 1994 the trial fishery was conducted with small Greenlandic boats in cooperation with a Norwegian mothership. L. maja and N. grimaldi were caught both with longline and gillnet, L. maja between 500-600 m and N. grimaldi between 800-900 m. In 1995 the mapping continued with two Norwegian gillnetters participating mainly as motherships, but also for mapping Greenland halibut (Reinhardtius hippoglossoides) distribution on the continental slope. Sub- stantial bycatch of L. maja had lead to the suggestion that this crab may be a potential resource on the east coast. One of the gillnetters, M/S Kato, therefore combined the gillnet fishery with pot fishery for L. maja. In 1996 the pot fishery continued with the Norwegian longliner M/S Skarheim in combination with the longline fishery in the fjords and on the shelf and continental slope. The objective of the trial pot-fishery survey was to evaluate the occurrence of L. maja according to area, depth, and temperature. Sex and size distribution, fecundity of females, and catch rates were investigated to gain a better understanding of the resource potential. Furthermore, processing of crabs, evaluation of meat quality, and market testing were done to evaluate the prospect for a high-quality and high-price product. Crabs in Cold Water Regions: Biology, Management, and Economics 735

Materials and Methods Study Area The trial fishery for L. maja in 1995 (August 14-September 6) was conducted in six different localities (Table 1; Fig. 1): between 62º and 63ºN on the continental slope (A, D), on the shelf (B, C, and E) and in the Ammasalik Fjord (F) at 65ºN. Fishing depth varied from 200 to 700 m. In 1996 (July 27-August 6) the fishery was conducted at about 62ºN in four different localities, from the continental slope (G) and westward on the shelf (H, I, and J) to the Tingmiarmiut Fjord (Table 1; Fig. 1). In general, localities in the continental slope were steep and rocky, while the bottom on the shelf was soft. Locality E was on the shallow shelf off the coast of Skjoldungen, an area dominated by cold water from the East Greenland current (Dietrich 1957). Locality A, B, G, H, I and J were off the coast of Tingmiarmiut Fjord, an area influenced more by the warmer Irminger Current (Dietrich 1957).

Gear The pots used were a modification of the model used in the Canadian snow crab fishery. They were conical, 67 cm high, with a top diameter of 65 cm and a bottom diameter of 120 cm. The pots were covered with 84 mm stretched mesh seine webbing. In 1995, the pots were baited with mackerel and squid, together weighing 0.5 kg. In 1996 an additional 0.2 kg of pollock were used. Soak time was planned to be 60 hours, but varied from 56 to 76 hours. In locality E, ice covering made hauling difficult and soak time was on average 105 hours. In Ammasalik Fjord, soak time was shorter (23 hours on average) in order to cover more of the fjord. Longlined strings of 30 crab pots (A-D), 15 pots (E-F), and 20 pots (G-J) were used. Pots were spaced 20 fathoms apart, giving mooring lines from 550 to 1,100 m.

Catch Data One station consisted of a string of crab pots. In 1995, a total of 38 stations were recorded (863 pots) harvesting 763 crabs (395 kg) (Tables 1 and 4). In 1996, a total of 12 stations were recorded (237 pots hauled) harvesting 1,134 crabs (460 kg) (Tables 1 and 4). Depth was classified in intervals of 100 m and fishing was conducted from interval 2 (200-300 m) to 6 (600-700 m) (Table 1). Evaluation of the bottom substrate was made from the echogram combined with observations on small samples of substrate picked up by pots and crabs. Tem- perature and salinity were measured with an electronic probe (SD202) from the surface to the bottom at the fishing localities (Table 3). Catch rates (CPUE) were estimated as weight (kg) of crabs per pot haul.

Crabs bigger than 0.6 kg were of commercial size (Brataas 1993). CPUEc was therefore estimated from the catch of crabs more than 0.6 kg. 736 Woll and Burmeister — Northern Stone Crab

Table 1. Position, depth interval, number of stations, and pots in the trial fishery for the northern stone crab (Lithodes maja) at southeast Greenland in August-September 1995 and July-August 1996.

Position Soak Depth a Latitude Longitude Stations Pots time intervals Locality (N) (W) (n) (n) (hrs) (100 m)

A slope 62.12-62.14 40.26-40.29 4 120 56 3-6 B shelf 62.25-62.29 40.48-40.58 8 240 65 2-5 C shelf 62.50-62.51 40.43-40.44 4 120 76 3-5 D slope 63.10-63.13 40.05-40.06 4 119 105 2-4 E shelf 63.10-63.12 40.12-40.30 6 882 65 2-3 F fjord 65.39-65.47 37.03-37.20 12 176 23 2-5

Total in 1995 38 863

G slope 62.09-62.13 40.29-40.37 4 80 66 4 H slope 62.10-62.11 40.37-40.39 2 40 60 3-4 I shelf 62.16-62.25 41.15-41.26 4 79 68 4-5 J shelf 62.33-62.35 41.59-42.01 2 38 24 4-5

Total in 1996 12 237 aThe number of pots varied between 15, 20, and 30 pots in each fleet.

Biological Data Carapace lengths (CL) were measured with calipers to the nearest millimeter. Live weight, sex, and presence of females with egg clutches were also recorded. A sample of 129 egg-carrying females was frozen for fecundity analyses. The eggs were later stored in 4% buffered formaldehyde. In the laboratory the egg clutch from each female was weighed and three subsamples of about 200 eggs each were randomly selected, then weighed and counted. Fecundity was determined by dividing the weight of the whole clutch by the average of the three estimates of individual egg weight. The mean egg diameter for one female (mean of length and width) was found by measur- ing 30 eggs to the nearest 0.1 mm with an ocular micrometer.

Processing and Market Testing Meat yield was tested regarding weight loss of whole cooked crabs in relation to live weight on six representative subsamples of 10-16 crabs each. For acceptable quality weight loss should be less than 25% (Brataas 1993). Crabs more than 0.6 kg were gutted as whole crab or clusters, and boiled for 20 minutes, after which the product was cooled, single frozen, glazed, and packed in boxes containing 5-6 kg of crabs. Only crabs with hard shells, indicating high meat yield, were used in the market testing. Be- cause of the small quantity of the product, the test was limited to restaurants in order to launch the crab as a high-quality and high-price product. Crabs in Cold Water Regions: Biology, Management, and Economics 737

Figure 1. The study area for the trial fisheries for the northern stone crab (Lithodes maja) in August-September 1995 (locations A to F) and July-August 1996 (locations G to J).

Results Catch Rates

The overall catch rate (CPUEtot) was 0.39 kg per pot in 1995 and 1.94 kg per pot in 1996 (Table 2). By weight, 61% of the crabs in 1995 were of commercial size (more than 0.6 kg) and 42% in 1996 (Table 2). The overall catch rate for crabs more than 0.6 kg (CPUEc) was 0.23 kg per pot (ranging from 0.04 to 0.47 among the localities) in 1995, and 0.81kg per pot (ranging from 0.19 to 1.43) in 1996 (Table 2). Catch rates for each depth interval from 200 to 700 m were estimated for all localities in 1995. The highest catch rate was achieved at depths between 400 to 500 m (Fig. 2), although there was no statistically significant difference between the catch rates at the five depth intervals (one way ANOVA, F(4,33) = 0.92, P = 0.46)

The highest CPUEc was achieved in 1996 on the continental slope at depths from 380 to 510 m, temperatures from 4º to 5ºC and on steep and rocky bottom at locality G and H (on average 1.43 kg per pot and 0.98 kg per pot, respectively) (Fig. 3; Table 3). 738 Woll and Burmeister — Northern Stone Crab

Table 2. Catch rates for Lithodes maja at southeast Greenland in 1995

and 1996. Catch rates expressed as total kg per pot (CPUEtot) and

as crabs more than 0.6 kg per pot (CPUEc).

All crabs Crabs more than 0.6 kg

Mean a Locality CPUEtot S.D. CPUEc S.D. (%) weight (kg)

A 0.53 0.61 0.47 0.54 90 0.97 B 0.68 0.55 0.39 0.37 57 0.83 C 0.45 0.19 0.18 0.06 40 0.76 D 0.55 0.24 0.39 0.15 70 0.89 E 0.38 0.42 0.17 0.19 45 0.75 F 0.09 0.15 0.04 0.08 44 0.72 1995 0.39 0.42 0.23 0.29 61 0.85

G 1.92 0.40 1.43 0.39 74 0.87 H 2.24 0.09 0.98 0.24 44 0.92 I 2.08 0.95 0.19 0.12 9 0.76 J 1.39 0.10 0.63 0.18 47 0.75 1996 1.94 0.61 0.81 0.59 42 0.81 a % of CPUEtot

Figure 2. CPUE (kg per pot) in different depth intervals during the trial fishery for the northern stone crab (Lithodes maja) in August-September 1995. Num- ber of pot hauls is marked over columns, number of stations in brackets. Crabs in Cold Water Regions: Biology, Management, and Economics 739

Figure 3. Catch rates of commercial sized crabs (CPUEc = kg crabs more than 0.6 kg per pot) during the trial fisheries for Lithodes maja in 1995 (locations A to F) and 1996 (locations G to J). Standard deviation marked with bars.

Table 3. Temperature recorded on the fishing grounds during the trial fishery for Lithodes maja in August-September 1995 (locations A, C, E, and F) and July-August 1996 (locations G, H, and J).

Temperature (ºC) Depth

(m) A C E F G I J

0 4.7 5.1 1.4 2.2 8.5 3.9 –0.1 50 4.7 5.1 0.6 0.1 7.6 4.4 –1.6 100 5.8 1.8 –0.4 –1.0 6.5 5.4 –1.6 200 5.4 3.0 0.0 –0.5 6.3 5.1 1.0 300 4.3 3.5 2.4 0.7 5.9 5.1 2.5 400 4.4 3.8 – 1.6 5.2 5.2 3.3 500 4.4 – – 1.7 5.0 4.9 3.8

600 – – – – 5.0 4.9 4.0

Temperatures at fishing depths are in bold. 740 Woll and Burmeister — Northern Stone Crab

The lowest catch rates were recorded in 1995 in Ammasalik Fjord (F) where 176 pot hauls gave a CPUEc of 0.04 kg per pot. Soak time for this locality was on average 23 hours compared to 56 to 105 hours in the other localities (Table 1). Bottom temperature varied from –0.5 to 1.6ºC. The bottom temperature at the other localities varied from 2.4 to 5.2ºC (Table 3).

Sex and Size Distribution Of the total number of crabs caught: 17.5% and 52.9% were females in 1995 and 1996, respectively. Sex ratio varied considerably among localities. On the continental slope just a few females were caught: none at locality A, 15% at locality D in 1995, and 7% at locality G in 1996. On the shelf, the catch of females varied from 13 to 21% in 1995 (locations B, C, and E) and from 50 to 75% in 1996 (locations I and J) (Table 4). The crabs and the pots at locality H, I, and J were partly covered with clay, indicating soft bottom. Size distribution of the females in 1995 showed that most of the individuals were between 70 and 75 mm CL (74 mm on average), varying from 45 to 106 mm. In 1996, most of the individuals were from 75 to 80 mm (77 mm on average), varying from 45 to 125 mm. (Fig. 4). For the males most of the crabs were from 85 to 110 mm CL (91 mm on average), varying from 37 to 134 mm CL in 1995. In 1996 the majority were between 85 to 110 mm CL (90 mm CL on average), varying from 47 to 126 mm CL. The length-weight relationship was established for females and males between 45 and 105 mm. It suggested that for crabs more than 60 mm CL, males were heavier than females of the same size in CL.

–3 2.879 2

Wfemales = 1.0 ¥ 10 ¥ CL r = 0.925; n = 131

–4 3.209 2 Wmales = 3.0 ¥ 10 ¥ CL r = 0.970; n = 628

Reproduction and Fecundity Approximately one-half of the females were egg carrying in 1995 and 1996 (Table 5). Females with egg clutches were bigger than 50 mm CL, indicating that this was close to the lower size at which they mature. The size of 50% maturity was 75 mm CL (Fig. 5). Some of the females had uneyed eggs and some had eyed eggs, indicating that hatching would soon happen. Some females had only remains of egg capsules on the pleopods, indicating they had just hatched. Fecundity varied from 2,000 eggs for females of 60 mm CL, to 15,000 for one female of 119 mm CL, and was positively correlated to CL (Fig. 6). Crabs in Cold Water Regions: Biology, Management, and Economics 741

Table 4. Sex ratio and mean carapace length (CL) for Lithodes maja recorded at southeast Greenland in August-September 1995 and July-August 1996.

Females Males Total

Locality (n) (% of n) CL (mm) S.D. (% of n) CL (mm) S.D.

A720––100 107 12 B 349 21 74 12 79 88 17 C 127 19 71 9 81 86 13 D 106 15 88 13 85 94 16 E701372628 88 11 F39266212388520 1995 763 17.4 74 13 82.6 91 17

G 228 7 80 11939713 H 176 40 90 11 60 91 14 I 603 75 75 8 25 78 16 J 127 50 76 10 50 91 15 1996 1134 52,9 77 10 47.1 90 16

The equation:

3.1186 2 Y = 0.0048 ¥ CL (R = 0.7405; n = 129), where Y = fecundity, describes the relationship between number of eggs and CL. Mean egg diameter of the individual females varied from 1.97 to 2.32, with a mean of 2.16 mm (S.D. = 0.06). The egg size was independent of CL.

Processing and Market Testing Crabs of commercial size (more than 0.6 kg) comprised 61% of the catch. These were boiled and frozen onboard. Weight loss for whole boiled crabs compared to live weight for representative subsamples from different localities varied from 24.7% to 36.2% (Table 6). This indicated that meat yield on average was low. Only hard-shelled crabs, indicating high meat yield, were used for the market test in the restaurants. The consignees were willing to pay more for live crabs (Norwegian Kroner, NOK 100-120 per kg) or fresh boiled crabs (NOK 90 per kg) compared to the frozen product they received. Handling of crabs was difficult because of the long spines. However, all the restaurants concluded that the meat was of excellent quality, comparable to the red king crab. The bright red coloration gave the crab an attractive appearance. 742 Woll and Burmeister — Northern Stone Crab

15 1995 n=763 10

5 Percentage of total

0 45-50 65-70 75-80 35-40 85-90 55-60 95-100 105-110 115-120 125-130

15 1996 n=1134 10

5 Percentage of total 0 45-50 65-70 75-80 85-90 35-40 55-60 95-100 Carapace length (mm) 105-110 115-120 125-130

female male

Figure 4. Size and sex distribution for catches of Lithodes maja at southeast Greenland in 1995 and 1996. n = number of crabs. Crabs in Cold Water Regions: Biology, Management, and Economics 743

Table 5. Egg-bearing females recorded in the catches of Lithodes maja at southeast Greenland in August-September 1995 and in July-Au- gust 1996.

1995 1996

Females Egg-bearing Females Egg-bearing

CL(mm) (n) (n) (%)a (%)b (n) (n) (%)a (%)b

45-50 2 0 0 0.0 1 0 0 0 50-55 3 1 33.3 1.5 7 0 0 0 55-60 10 6 60.0 10.6 11 1 9.1 0.3 60-65 15 4 26.7 16.7 27 8 29.6 3.0 65-70 18 6 33.3 25.8 83 35 42.2 14.4 70-75 30 13 43.3 45.5 108 54 50.0 32.1 75-80 21 8 38.1 57.6 126 66 52.4 53.8 80-85 7 5 71.4 65.2 105 67 63.8 75.7 85-90 7 6 85.7 74.2 63 37 58.7 87.9 90-95 11 9 81.8 87.9 32 18 56.3 93.8 95-100 6 6 100 97.0 18 11 61.1 87.4 100-105 1 0 0 97.0 10 6 60.0 99.3 105-110 2 2 100 100 1 1 100 99.7 110-115 0 99.7 115-120 0 99.7 120-125 1 1 100 100

Total 133 66 49.6 593 305 51.4 a% of females in the CL interval. b% cumulative of egg bearing females. 744 Woll and Burmeister — Northern Stone Crab

Figure 5. Size distribution for females with and without egg clutches in 1995 (n = 133) and 1996 (n = 593).

Table 6. Weight loss for boiled crabs compared to live weight for Lithodes maja on the southeast coast of Greenland in Au- gust 1995 (locations B, C, D, and E) and 1996 (locations G and I).

Mean Loss of

Crabs weight (kg), weight (%),

Locality (n) live boiled

B160.65 24.7 C100.66 34.9 D101.06 31.9 E120.67 30.8 G100.87 35.0 I100.84 36.2 Crabs in Cold Water Regions: Biology, Management, and Economics 745

Figure 6. Estimated fecundity for females of Lithodes maja from a representative sample in 1996.

Discussion Catch Rates Fishing trials in 1995 were conducted over a wide area, in depth intervals from 200 to 700 m and bottom temperature ranging from –0.5º to 4.4ºC. The presumption undertaken after the fishery in 1995, that the bigger males were more abundant near the continental slope on rocky bottom, was verified by the results in 1996. Both years, the highest catch rates of commercial sized crab were achieved at such localities. Females were more common on soft bottom, often together with smaller crabs of both sexes. In 1996 fishing depths were narrowed to 350-500 m where bottom temperatures were 3.3º to 5.2ºC. This resulted in higher catch rates (CPUEc = 0.81 kg per pot). Several trial fisheries for L. maja have been conducted along the south coast of Newfoundland. In October 1992 a CPUE of 4.0 kg per pot was obtained on depths from 150 to 250 m on hard, rocky bottom (Dooley and Johnson 1993). From August 1994 to March 1995 CPUE was on average 1.1 kg per pot (48 hours soak time) (DFOC 1995) and from September to De- cember 1996, 1.3 kg per pot (CAFID 1996). Crabs more than 95 mm carapace width (CW) are considered market size in Canada (CAFID 1996), and only crabs of this size were kept. CL and CW are nearly the same for L. maja (Andersen 1993, Woll 1996), and 95 mm CL corresponds to approximately 0.65 kg for male crabs. The catch rates achieved in southeast 746 Woll and Burmeister — Northern Stone Crab

Greenland were, according to this size to weight relation, lower than the catch rates achieved in Newfoundland. The trial fishery at southeast Greenland was conducted in August. In northern Norway, a trial fishery was conducted from October 1992 to March 1993. The catch rates decreased in March, suggesting that the crab had seasonal migrations and that catchability changed, probably due to molting (Hufthammer 1996). The highest catch rates in Norway were obtained at depths from 100 m to 200 m. The females were most abundant from 150 to 200 m and the males at depths from 100 to 200 m (Hufthammer 1996). The difference in abundance at different depths may be due to seasonal effects as crabs usually have seasonal migrations. A fishery at southeast Greenland later in the autumn may give different catch rates due to this. Bottom temperature is probably of importance for the presence of crabs. The best catches at southeast Greenland were obtained at about 4 to 5ºC. No temperature data were reported from Norway and Newfound- land for comparison. In 1996 a few strings of pots were set near the Tingmiarmiut Fjord. The catch rate of commercial crabs was lower than on the slope, but higher than on the shelf. No pots were set in Tingmiarmiut Fjord, but two big males were caught on a longline. Ice covering during the trial fishery in 1996 prevented further exploration of Tingmarmuiut Fjord (Woll and Gundersen 1997). However, in summer 1998 L. maja was caught as bycatch on small meshed gillnets in Lindenow Fjord at depths from 350 to 400 m (Woll et al. 2000).

Reproduction and Fecundity Egg size for L. maja at southeast Greenland was on average 2.16 mm. Reported egg size for L. maja in Scotland (Clyde area) was slightly smaller, about 1.8 ¥ 2.0 mm when mature (MacDonald et al. 1957). The egg size of L. maja is large compared to Paralithodes spp. This may be due to the fact that all larval stages of L. maja, from hatching to metamorphosis, are lecithotrophic (Anger 1996). This is considered an adaptation to seasonally short and limited planktonic food production in subarctic regions of the northern Atlantic (Anger 1996). The presence of egg capsules on the pleopods of the females indicates that hatching occurred for a part of the females in July to August at southeast Greenland. The presence of uneyed and eyed eggs indicates that hatching probably takes place over a longer period, which is possibly due to the lecithotrophic larvae. Somerton and Otto (1986) concluded that the related Lithodes aequispinus appear to have protracted, or perhaps year-round, breeding due to the lack of a clear seasonal change in the occurrence of eyed and uneyed clutches. An alternative strategy to protracted breeding is for reproduction to be synchronized with climatic cycles, so that larval hatching occurs only during a short period in spring, at the beginning of a seasonal plankton bloom. This is found in other subarctic Crabs in Cold Water Regions: Biology, Management, and Economics 747 species, e.g., in the red king crab Paralithodes camtschaticus (Dawson 1989). MacDonald et al. (1957) reported that in the Clyde area (U.K.), spawning occurred in autumn (September to November) and females carried the eggs throughout the winter months. Hatching took place in April or May and the parent molted soon afterward in preparation for a second brood.

Processing and Meat Yield Meat yield for crabs caught at southeast Greenland in August was low compared to meat yield during the trial fishery in Norway in October (Brataas 1993). In southeast Greenland some of the crabs were soft shelled indicating that molting occurs in summer for parts of the stock. The trial fishery in Canada and Norway took place during October to March, and meat yield seemed to be good at this time of the year. According to this, meat yield for L. maja at southeast Greenland may be higher in autumn.

Conclusions L. maja is abundant on the slope, the shelf, and in the fjords at southeast Greenland. Males are larger than females, and approximately all crabs of commercial size (more than 0.6 kg) were males. In August large males seemed to be most abundant on the continental slope at depths from 300 to 500 m, temperatures of 4º to 5ºC and on steep, rocky bottom. The highest CPUEc obtained on such a locality was 1.43 kg per pot. Catch rates were on average smaller than obtained in a trial fishery in Newfoundland where the favorable fishing season seems to be from October to January. At this time of the year, fishing is complicated at southeast Greenland because of ice cover and bad weather. The limited catch data makes an evaluation of the prospect for a profitable fishery for L. maja at southeast Greenland difficult, and further investigation of catches by season must be undertaken to get a better understanding of this. However, the catch rates and estimated prices are too low to give more backing to this project at present.

Acknowledgment The project was financed by the Norwegian Research Council, the Nordic Council of Ministers, Greenland Home Rule, and Greenland Institute of Natural Resources. The skippers and crew on board M/S Kato in 1995 and M/S Skarheim in 1996 did a good job. Technician Kunuk Kloster from Greenland Institute of Natural Resources assisted during the survey in 1995, and in 1996 together with technician Jan Erich Rønneberg from Møre Research. Margaret Kjerstad, Møre Research, was responsible for the market testing together with three Norwegian companies: North Cape Fish, Seafood, and Ålesundfisk. Agnes C. Gundersen and Stig Tuene provided good advice to improve the manuscript. A big thank you to all of them. 748 Woll and Burmeister — Northern Stone Crab

References Anger, A. 1996. Physiological and biochemical changes during lecithotrophic larval development and early juvenile growth in the northern stone crab Lithodes maja (Decapoda: Anomura). Mar. Biol. 126:283-296. Andersen, M. 1993. Krabber og krabbefiseri ved Vestgrønland. Grønlands Fiskeri- undersøkelser, Nuuk. 50 pp. Bratteng, A. 1994. Markedsføring av trollkrabbe (Lithodes maja) i Tyskland. Norwe- gian Seafood Export Council, Tromsø. 22 pp. Brataas, R. 1993. Prøveproduksjon og markedsvurdering av trollkrabbe (Lithodes maja). Norwegian Institute of Fisheries and Aquaculture, Tromsø. 14 pp. CAFID. 1996. Development of the Atlantic king crab fishery. Project summary. Can- ada/Newfoundland Cooperation Agreement for Fishing Industry Development (CAFID), No. 21. 4 pp. Dawson, E.W. 1989. King crabs of the world (Crustacea: Lithodidae) and their fisheries: A comprehensive bibliography. New Zealand Oceanogr. Inst., Div. Water Sci, DSIR (Department of Scientific and Industrial Research) Misc. Publ. 101, Wellington. DFOC. 1995. Northern stone crab. Coordinator/Monitor Project Summary Report. Department of Fisheries and Oceans Canada (DFOC), Industry Development Division. 31 pp. Dietrich, G. 1957. Schictung und Zirkulation der Irminger Sea im Juni 1955. Ber. Dtsch. Komm. Meeresforsch. 14:255-312. Dooley, T., and B. Johnsen. 1993. Exploratory fishing spiny crab, south coast 1992. Government of Newfoundland and Labrador, Department of Fisheries, Har- vesting and Operation Division. 7 pp. Hufthammer, M.K. 1996. Utbreiing, mengde, bestandsstruktur, reproduksjon og ernæring hos trollkrabbe (Lithodes maja). Hovedfagsoppgave i fiskeribiologi, University of Bergen, Department of Fisheries and Marine Biology. 53 pp. MacDonald, J.D., R.B. Pike, and D.I. Williamson. 1957. Larvae of the British species of Diogenes, Pagurus, Anapagurus and Lithodes (Crustacea, Decapoda). Proc. Zool. Soc. London 128:209-257. Somerton, D.A., and R.S. Otto. 1986. Distribution and reproductive biology of the golden king crab, Lithodes aequispina, in the eastern Bering sea. Fish. Bull., U.S. 84(3):571-582. Squires, H.J. 1990. Decapod Crustacea of the Atlantic Coast of Canada. Can. Bull. Fish. Aquat. Sci. 221. 323 pp. Woll, A. 1996. Exploratory fishery for the northern stone crab (Lithodes maja) off the southeast coast of Greenland. ICES C.M. 1996/K:19. Woll, A.K., and A.C. Gundersen. 1997. Kartlegging av fjorder og kontinentalsokkel ved Sydøst-Grønland. Topografi, hydrografi og fiskeressurser. Møre Research, Å9719, Ålesund, Norway. 66 pp. Crabs in Cold Water Regions: Biology, Management, and Economics 749

Woll, A.K., E. Hjörleifsson, and J. Boje. 2000. Exploratory fishery for young Green- land halibut around southern Greenland. M/S Aud-Lill 1998. In: Greenland halibut in east Greenland waters. Nordic Council of Ministers, TemaNord Fisheries 2000(585):113-136.

Crabs in Cold Water Regions: Biology, Management, and Economics 751 Alaska Sea Grant College Program • AK-SG-02-01, 2002

Review of the Family Lithodidae (Crustacea: Anomura: Paguroidea): Distribution, Biology, and Fisheries

S.D. Zaklan University of Alberta, Biological Sciences, Edmonton, Alberta, Canada

Abstract The family Lithodidae is a diverse group of decapods on which investigative research is coming of age. Herein, published literature and original data are compiled and summarized. This overview includes distributions, life history parameters, morphology, and names. Lithodids are a large family of approximately 105 species that are broad-scale omnivores with seasonal reproduction. They have a global distribution but reside mainly in antitropical waters from intertidal to 4,152 m. They most likely arose in the North Pa- cific Ocean 13-25 million years ago. Much remains unknown about this economically important group, most likely due to their abyssal nature. Before more crab fisheries collapse or new fisheries are opened, increased research and enhanced communication among crab biologists should be initiated.

Introduction Deep-sea members of the family Lithodidae Samouelle, 1819 (Crustacea: Decapoda: Anomura) rank among the world’s largest arthropods. The family Lithodidae is divided into two subfamilies (Hapalogastrinae and Lithodinae Ortmann, 1901) that collectively include 15 (Sakai 1976) or 16 genera (Dawson 1989) and approximately 105 species (Table 1) that reside mainly in the North Pacific Ocean (Table 2). Lithodid characteristics include a crab-like exoskeleton, rudimentary fifth walking legs, no uropods, asymmetric abdominal tergite plates, and pleopods 3-5 found only on the left. However, the majority of lithodid characters are based on their unusual abdomen. First, the sternal plate of the first abdominal segment is articulated with the last thoracic segment. Second, the female’s abdomen is distinctly asymmetrical, with the medial plane of symmetry right of 752 Zaklan — Review of Lithodidae center. Third, while males do not have pleopods, females possess between four and six asymmetrically placed pleopods, with the majority (four or five) associated with her larger left abdominal plates. Finally, there are varying degrees of abdominal tergal plate calcification, from none (Oedignathus) to complete (e.g., Cryptolithodes; e.g., Makarov 1962, Rich- ter and Scholtz 1994). These unusual morphologies associated with asymmetry and variable calcification have puzzled evolutionary biologists for over one hundred years (Boas 1880a,b; Bouvier 1894a,b, 1895b,c, 1896, 1897; Borradaile 1916), and recently there has been a resurgence of interest in lithodid evolution (Cunningham et al. 1992; Richter and Scholtz 1994; McLaughlin and Lemaitre 1997, 2000; Zaklan 2001).

Identity and Relationships Lithodid evolutionary relationships were first proposed by Boas (1880a,b) and Bouvier (1894a,b, 1897), who suggested that the asymmetrical soft abdomen of king crabs evolved from the asymmetrical soft abdomen of an ancestor that resembled hermit crabs and inhabited gastropod shells. Recent molecular (Cunningham et al. 1992, Zaklan 2001; Fig. 1) and some (Richter and Scholtz 1994) but not all (Martin and Abele 1986; McLaughlin and Lemaitre 1997, 2000) morphological reconstructions agree with this hypothesis. Broad scale lithodid genus-level relationships, based on morphology of the abdomen, were first hypothesized by Bouvier (1897) and recently confirmed by the molecular phylogeny found in Zaklan (2001; Fig. 1). Although there is no single identification manual that distinguishes all known lithodid species, the most inclusive keys are found in Dawson and Yaldwyn (1985a) and Macpherson (1988c). Other helpful keys are referred to in Table 3. Original taxonomic descriptions (Table 1) also may be used to differentiate potentially confusing species (Table 4). The Web site http://geocities.com/Lithodidae is useful for viewing photographs and accessing reference information.

Origin and Distribution Lithodids are distributed across a variety of aquatic zones from mid-intertidal to abyssal depths (to 4,152 m for Paralithodes bouvieri, Macpherson 1988c). They typically inhabit anti-tropical waters, with high concentra- tions in the northern Pacific Ocean (Table 1). Lithodid radiation most likely began in the intertidal zone of the northeastern Pacific (Bouvier 1896, Makarov 1962) from a hermit crab–like ancestor (Boas 1880a,b; Bouvier 1894a, 1897; Cunningham et al. 1992; Richter and Scholtz 1994; Zaklan 2001; but see McLaughlin and Lemaitre 1997, 2000). The family is of fairly recent origin, arising between 13-25 mya as suggested by molecular (Cunningham et al. 1992) and fossil (Feldmann 1998) evidence, and possibly evolving under the protection of the recently evolved (16-30 mya; Estes and Steinberg 1988, Saunders and Druehl 1992) canopy-producing kelp order Laminariales (Zaklan 2001). Crabs in Cold Water Regions: Biology, Management, and Economics 753

Makarov (1962) provided several lines of biogeographic evidence that suggested a northeastern Pacific Ocean lithodid origin. Recent observations (Tables 1 and 2) concur with Makarov’s North Pacific hypothesis. First, 68% of species and 100% of genera are reported from the Pacific, compared to only 28% of species and 19% of genera from the Atlantic Ocean. The remaining species are found in the Indian (11%) and Antarctic oceans (3%; note: since species have broad distribution ranges, locality percentages do not add up to 100%). Second, 55% of living lithodid species occur in the Northern Hemisphere, as compared to only 42% in the south, the remaining taxa being abyssal inhabitants of the tropics (Tables 1 and 2). Third, the subfamily Hapalogastrinae is basal within the family Lithodidae (Bouvier 1894b, 1897 as morphological evidence; Konishi 1986 as larval evidence; Zaklan 2001 as molecular evidence; see Fig. 1). This basal subfamily is found only in the North Pacific Ocean, mainly in the northeastern Pacific (89% of species compared to only 45% in the northwestern Pacific, Tables 1 and 2). Finally, two basal clades (Fig. 1; according to Zaklan 2001) within the Lithodidae, Hapalogastrinae and genus Cryptolithodes of the Lithodinae, occur in the intertidal or shallow subtidal zone, suggesting an intertidal origin. Although most intertidal lithodids inhabit the northeast Pacific, six intertidal species exist outside this region, five of which reside in the northwest Pacific. The sixth intertidal inhabitant is Paralomis granulosa, a shallow-water inhabitant, that apparently recently colonized southern South America’s Beagle Channel, after the last deglaciation (8,200 years ago; Makarov 1962, Rabassa et al. 1986). This is the only member of the genus that inhabits shallow coastal waters (Macpherson 1988). This species also retains certain reproductive features associated with varying or unpredictable food supplies such as few large eggs and a reproductive cycle in which zoeal eclosion is independent of food supply (Lovrich and Vinuesa 1993, 1999). From the northeast Pacific’s intertidal, lithodids are thought to have advanced west along the Aleutian range to the northwest Pacific shores of east Kamchatka and the Kuril Islands of Russia, followed by a range expansion to eastern Asia’s Japan, Korea, and China. Concurrently, northeast Pacific lithodids spread south along the eastern Pacific shores of North America. Due to cold-water abyssal upwelling, intertidal species are found farther south (Baja California) than their northwestern Pacific counterparts (Japan, Makarov 1962; Table 1). Deep-sea lithodids probably crossed the tropics into the temperate, subantarctic, and antarctic regions of the South- ern Hemisphere where a limited recolonization of South America’s intertidal occurred (Makarov 1962, Lovrich and Vinuesa 1993; Table 1). Lithodids presumably passed through the Antarctic Ocean and spread upward through to the Atlantic, across the southern tip of Africa into the Indian Ocean, and finally into the southeast Pacific (Makarov 1962). Along with the Pacific radiation of lithodids was a northern Atlantic radiation. Lithodids are thought to have passed north from the northeastern 754 Zaklan — Review of Lithodidae

Pacific by way of the Bering Sea through the Arctic Ocean, spreading throughout the North Atlantic and southward around Africa into the In- dian Ocean. Lithodids found in the Indian ocean are thought to have moved by way of the Atlantic Ocean and not via the western Pacific Ocean due to the lack of observed lithodid inhabitants in the waters surrounding western Australia and Indonesia (p. 35, Fig. 12 in Makarov 1962; for an overview of their present distribution see Table 1). Perhaps the abyssal nature of these animals (depth averaged from Table 1 minimum and maximum depths: Neolithodes = 1,570 m; Lithodes = 532 m; Paralomis = 821 m; Table 1), along with a large protective exoskeleton (CW £ 300 mm, Table 4), group social dynamics, migratory abilities, large broods (up to 280,000 eggs; Matsuura et al. 1972), expansive larval dispersal capabilities (Table 5) and opportunistic foraging strategies, allowed them to spread into extreme habitats that offer little environmental protection from overhead predators (Table 6).

Sperm Morphology Only three lithodid species have been examined for spermatozoal morphology: one hapalogastrine, Hapalogaster dentata (Goshima et al. 1995); and two lithodines, Lithodes maja (Retzius 1909, Tudge et al. 1998) and Paralithodes camtschaticus (Marukawa 1933). Overall, their spermatozoa are spherical, and have globular nuclei and concentrically zoned acrosome vesicles. They are topped by a centrally perforated operculum, and are penetrated by a perforatorial chamber that is posteriorly embedded in the cytoplasm (Jamieson and Tudge 2000:32). The spermatophores and sperm of L. maja share many synapomorphic features with the hermit crab genus Pagurus. These include accessory ampulae, homogenous granular spermatophore wall, concentric zonation of the acrosomal vesicle, operculum shape and differentiation, and an electron-dense plume basally in the perforatorial chamber (Tudge et al. 1998).

Parasites There are three major types of lithodid parasites: rhizocephalans (Briarosaccus spp.), liparids (Careproctus spp.), and microsporidians (Table 7). Briarosaccus callosus (Cirripedia: Peltogastridae) has four naupliar stages and one cyprid stage. It is the primary rhizocephalan barnacle parasite as it is hosted by several lithodids and has a nearly worldwide distribution (Boschma 1970; Table 7). Briarosaccus tenellus has five naupliar stages and has been found associated only with Hapalogaster mertensii (Boschma 1970, Walossek et al. 1996). Larval morphology indicates that rhizocephalans hosted by Paralithodes camtschaticus, P. platypus, and L. aequispinus are conspecifics (Hawkes et al. 1985a), and hemolymph responses and electrophoresis lend additional support to this hypothesis (Shirley et al. 1986). Rhizocephalans cause feminization through castra- tion and reduce growth in both sexes. Parasitized P. platypus and L. Crabs in Cold Water Regions: Biology, Management, and Economics 755 aequispinus are smaller than their unparasitized conspecifics (Sloan 1984, Hawkes et al. 1986a, 1987) and lithodids with multiple infections (up to five, observed in fjord dwelling northern British Columbia Lithodes aequispinus) are even smaller (Sloan 1984). Parasitism can potentially affect reproductive stock as infection levels of B. callosus can range from 40.5% to 66.7% of the total population (McMullen and Yoshihara 1970, Sloan 1984). However, occurrence is typically less than 1% in commercial landings of legal P. camtschaticus and L. aequispinus and up to 12% in P. platypus (Hawkes et al. 1986b). The other major lithodid parasite is the liparid fish genus Careproctus. Some species in this snailfish genus oviposit their eggs into the protected cavity of aerated lithodid gill chambers. Although Careproctus eggs are found in relatively few species of king crabs (Table 7) they can exist in up to 43.6% of the population (L. aequispinus; Jewett et al. 1985, Love and Shirley 1993, Somerton and Donaldson 1998). Negative effects of parasitism include egg mass induced gill compression (Anderson and Cailliet 1974, Melville-Smith and Louw 1987, Somerton and Donaldson 1998), gill bleeding (Love and Shirley 1993), and gill necrosis. In extreme cases gills are reduced to blackened stubs and up to 35% mortality is observed (Somerton and Donaldson 1998). Nemertean brood symbionts such as Carcinonemertes regicides and Alaxinus oclairi also are thought to cause reductions in P. camtschaticus populations by eliminating recruitment of some year classes to the fishery through brood mortality (Gibson et al. 1990, Kuris et al. 1991). Alaxinus oclairi can reduce broods by more than 50% (Gibson et al. 1990), and C. regicides were thought to be responsible for near complete consumption of all brooded eggs in some localities during the P. camtschaticus 1983- 1985 breeding season (Kuris et al. 1991). Other egg predators such as amphipods as well as viral infections and microsporidans may have an impact on brood success and population growth (Sparks and Morado 1985, Kuris et al. 1991; Table 7).

Subfamilies Subfamily Hapalogastrinae This taxonomically small subfamily of nine species (if Placetron forcipatus is recognized as a species; see Dawson 1989) and five genera is found only in the northern Pacific Ocean, mainly in the northeast (89% of species; Table 2). Species inhabit intertidal and shallow subtidal (down to 245 m for Acantholithodes hispidus [United States National Museum of Natural History; U.S.N.M.]) rocky shores and reside in a variety of protective habitats such as rocks, crevices and kelp (order Laminariales). They are generally opportunistic suspension feeders (Table 6). Relatively little natural history is known about this group, except that they are annual spawners that lack any seasonal- or size-dependent movement patterns (Goshima et 756 Zaklan — Review of Lithodidae al. 1995; Table 5). Although some members of genus Hapalogaster can be found in groups (Goshima et al. 1995), mass gatherings or Paralithodes camtschaticus–like “pods” (see below) have never been documented. Mem- bers of Oedignathus inermis are found only in pairs during the August mating season, and they remain solitary dwellers of crevices for the remainder of the year (Zaklan, pers. obs.). All Hapalogastrinae have four zoeal stages and one glaucothoe stage, whereas species in the subfamily Lithodinae have two to four zoeal stages. Lithodinae larval stage abbrevia- tion may suggest hapalogastrines are a basal lineage within the family Lithodidae (Konishi 1986).

Subfamily Lithodinae This speciose subfamily (96 species representing 10 genera if Acantholithus is a true genus; see Sakai 1976 and Dawson 1989) is globally distributed but is concentrated in benthic North Pacific waters. Although they are generally found between 100 and 1,000 m subtidally, they range in habitat from the low intertidal (Cryptolithodes; Hart 1965) to abyssal depths (Paralomis 4,152 m; Macpherson 1988c). All genera have North Pacific Ocean representatives whereas only 30% of genera are represented in other oceans (e.g., Atlantic, Antarctic, Arctic and Indian; Table 1). Many species are recorded only by their holotype or a single sex (e.g., Lithodes richeri, Macpherson 1990; L. wiracocha, Haig 1974; Paralomis zealandica, Dawson and Yaldwyn 1971; P. longidactyla, Birstein and Vinogradov 1972; P. microps, Filhol 1884; P. anamerae, P. erinacea, P. grossmani, P. pectinata, P. serrata, Macpherson 1988c; P. tuberipes, Macpherson 1988b; P. sp., Macpherson 1990; P. jamsteci, Takeda and Hashimoto 1990; Table 4). Thus, taxonomic, systematic, and distributional understanding of lithodines is still in its infancy. Species in this subfamily are known to have seasonal- and size-dependent movements, seasonal or aseasonal reproductive patterns, and omnivorous opportunist feeding habits (Table 6). Members of the subfamily Lithodinae generally have a greater and more abyssal geographic distribution, are more speciose (Table 1), are larger (Table 4), carry more eggs, have fewer zoeal stages (Table 5), and live in larger social groups than members of the subfamily Hapalogastrinae.

Subfamily Lithodinae Life Cycle The most notable characteristic of the Lithodinae is their migratory pattern, which is inextricably intertwined with mating rituals and with associated seasonal abiotic factors (Marukawa 1930, Bright 1967). Bathymetric location, movement patterns, and mass gatherings (podding) are influenced by increased benthic production (Rodin 1970, Stinson 1975), thermocline (Somerton 1985), halocline, timing of larval release, thermohaline mixing, temperature, food sources, and photoperiod in adults (Stone et al. 1992). Larval movement patterns are modified by temperature, light and salinity (Shirley and Shirley 1988, 1989b), as larvae are negatively geotactic and positively rheotactic (Shirley and Shirley 1988). King crabs are capable of Crabs in Cold Water Regions: Biology, Management, and Economics 757 long-distance navigation using both chemosensory cues and home-range environmental features as guides (Dew 1990, Stone et al. 1992). Substantial movements are a major part of their lives. They move from one area to another in discrete groups that are often segregated into sex and size-classes (Marukawa 1930, Powell and Nickerson 1965a, Bright 1967 for Paralithodes camtschaticus; Abello and Macpherson 1986 for L. ferox; Miquel et al. 1985 for L. murrayi). For example, female P. camtschaticus have a 3.6-11.9 km2 range (Stone et al. 1992) and males can move 112 km in 113 days in Alaska (Simpson and Shippen 1968). In Japan they can move up to 10.4 (females) and 13.1 (males) km per day (Marukawa 1933). Tracking benthic crustaceans has become easier with the advent of internal tags that can be retained throughout ecdysis, such as coded wire tags and passive integrated transponder tags (Donaldson 1997). Laser line scans (Tracey et al. 1998), submersibles (Zhou and Shirley 1998), and ultrasonic biotelometry (Stone et al. 1992) also can be used for identification and tracking purposes. Size at reproductive maturity is well documented for economically important lithodines compared to their unharvested conspecifics. Gener- ally, maturity is defined as gonadal maturity unless otherwise stated, but several measures of reproductive maturity exist. For example, crabs of a given size class are considered functionally mature when they have been observed procreating. Males are gonadally mature when they possess spermatozoa in deferent ducts, while females are considered gonadally mature when they have eggs attached to their pleopods (Lovrich and Vinuesa 1999). Members of the family Lithodidae are morphometrically mature when the relative growth of the right chela’s height changes with respect to carapace growth (Somerton and MacIntosh 1983). As size at female reproductive maturity is assumed to be the smallest ovigerous female documented or observed, carapace length at reproductive onset will decrease with future research and further published observations (Table 5). The majority of studied lithodines have a distinct, annual life history pattern associated with their reproductive cycle. Generally, females molt under the protection of a larger male. Molting is then followed by mating (Marukawa 1933, Powell and Nickerson 1965b, Somerton and MacIntosh 1983, Paul and Paul 1990, Vinuesa 1991, Goshima et al. 1995, Wada et al. 1997, Lovrich and Vinuesa 1999, Paul and Paul 2001b, Hoyt et al. 2002). They then carry eggs for approximately one year (Table 5), and if the zoeae are planktivorous, eclosion is concurrent with the spring diatom bloom (Kurata 1959, Paul and Paul 1980, Shirley and Shirley 1989a). Movement occurs throughout this cycle for all developmental stages, from vertical zoeal movements (Shirley and Shirley 1987, 1989b) to hori- zontally adult movements in genera such as Lithodes, Paralithodes, and Paralomis (i.e., Miquel et al. 1985, Vinuesa 1991, Abe 1992, Stone et al. 1992, Hoggarth 1993). Large-scale movements are unknown but probable for the long-legged Neolithodes, and unlikely for the more robust Phyllolithodes, Cryptolithodes, Rhinolithodes , and Lopholithodes. There are both inter- and intraspecific variations to this life history pattern. For example, life 758 Zaklan — Review of Lithodidae history patterns of Lithodes aequispinus are locality dependent (Rodin 1970) so that they are aseasonal (have members of the population in varying stages of their life cycle throughout the year) and migratory in fjords of British Columbia, Canada (Sloan 1985, Somerton and Otto 1986) whereas they are nonmigratory and reproduce seasonally on Japan’s continental shelf (Hiramoto and Sato 1970). Bottom temperature has a great effect on growth (Stevens 1990), and thus on lithodid fecundity. For example, extreme cold-water inhabitants, such as Paralomis granulosa (Lovrich 1997) and Paralithodes camtschaticus (McCaughran and Powell 1977, Zheng et al. 1995), take between 5 and 10 years to reach gonadal maturity as their molt interval is longer and their molt increment is less at colder temperatures (summarized by Loher et al. 2001). Differences in water temperature are thought to cause the inverse correlation (Jewett et al. 1985) between size at maturity and latitude in L. aequispinus found in the eastern Bering Sea (Somerton and Otto 1986) and P. camtschaticus. For example, female P. camtschaticus reach maturity at an average carapace width (CW) of 89 mm in Bristol Bay, but at 71 mm CW in Norton Sound (Fishery Management Plan for Bering Sea/Aleutian Is- lands King and Tanner Crabs 1998, North Pacific Fishery Management Council, Table E. 5).

Subfamily Lithodinae Behaviors All lithodines probably have anti-predator behavior; however, this is mainly documented in the economically important red king crab, P. camtschaticus. Following juvenile metamorphosis, these crabs are solitary and use crypsis and protected refuges afforded by complex habitats (e.g., rocks and kelp) in the intertidal and shallow subtidal as a sanctuary from benthic predators (Loher and Armstrong 2000). As they increase in size, they are often found in the protective crevices between starfish arms located on barnacle encrusted dock pilings (Evasterias troschelii and Asterias amurensis in Alaska; Powell and Nickerson 1965a, Dew 1990); and Leptasterias sp. in the Gulf of Shelikov, Russia; Vinogradov 1968). They are also found among Metridium senile anemone stalks, where they feed upon food particles dislodged by their commensal hosts (Powell and Nickerson 1965a). Juve- nile king crabs leave their protective niches after sunset and move into the open to forage (Dew 1990). The most striking behavior pattern of lithodids is “podding,” exemplified by both adult and juvenile P. camtschaticus. Podding describes a dense aggregation of hundreds to thou- sands or even to millions of similarly sized P. camtschaticus. Podding is thought to serve as a predatory-defense mechanism, and may also divide pre-mating adult populations into subgroups (Powell and Nickerson 1965a,b; Bright 1967; Dew 1990; Stone et al. 1993). Crabs in Cold Water Regions: Biology, Management, and Economics 759

Subfamily Lithodinae Adaptations Lithodines possess several adaptations that allow them to thrive in the vast open regions of the deep sea (Somerton 1981). First, some lithodids (Paralithodes, Lithodes, Neolithodes, Glyptolithodes, and some species of Paralomis) have long and slender pereiopods. The associated reduced musculature is considered less costly to maintain, and long legs allow rapid and efficient movement by taking fewer, larger steps during long distance movements (Somerton 1981). Second, L. couesi, the scarlet king crab, possesses red coloration that is found in many deepwater crustaceans and believed to represent cryptic coloration due to the rapid attenu- ation of red surface light (Marshall 1954). Third, deepwater inhabitants possess enlarged exhalent openings and scaphognathites, allowing for greater water volume to be pumped over gills compared to shallow water conspecifics (Somerton 1981). These enlarged gills may be associated with inflated branchial chambers, as found in Paralithodes camtschaticus, L. aequispinus, and Paralomis verrilli (Takeshita et al. 1978). Fourth, deepwater crabs such as Lithodes couesi and L. aequispinus have asynchronous or protracted spawning and breeding periods that are most likely a function of aseasonal productivity at great depths (Somerton 1981, Sloan 1985). Fifth, lecithotrophy and associated large eggs are adaptive for the vari- ably productive habitats of L. aequispinus (Shirley and Zhou 1997) and L. santolla (Lovrich 1999). Lecithotrophic development allows for successful hatching and recruitment, independently of phytoplankton blooms, thus allowing reproduction and subsequent larval release to be asynchronous (Comoglio and Vinuesa 1991, Shirley and Zhou 1997). Finally, L. couesi often live on remote seamounts on the continental slope that are isolated from the ocean surface, coastal areas, and possibly predators. These submarine islands can be inhabited by species that are able to tolerate vast ranges in depth and possess sufficient dispersal capabilities to move through the continental slope (Sakai 1971, Somerton 1981) or by species with larvae possessing adaptations that allow them to remain as seamount inhabitants. For example, lecithotrophic larvae of L. aequispinus remain near the bottom in laboratory cultures (Jewett et al. 1985, Shirley and Zhou 1997) and have never been collected in the plankton (Somerton and Otto 1986, Shirley and Shirley 1989a). These behavioral adaptations would limit their distribution and effectively confine lecithotrophic larvae within restricted habitats such as seamounts (T. Shirley, pers. comm.)

Subfamily Lithodinae Fisheries The Pacific Ocean contains extensive and diverse coastlines and open shelf areas that support some of the most commercially significant crustacean stocks in the world, including four species of king crab (Paralithodes camtschaticus, P. platypus, L. aequispinus, and L. couesi) in the north and two species in the south (Paralomis granulosa and L. santolla). The Bering 760 Zaklan — Review of Lithodidae

Sea, the Sea of Japan, the Kamchatka region, and the Gulf of Alaska were part of a rapidly expanding king crab fishery from 1960 to 1980 (Fig. 2). Peak landings of 84,000 t occurred in this male-only fishery in 1980. At this stage Paralithodes camtschaticus was the most valuable single-species fishery in the United States (168.7 million pounds; Alaska Department of Fish and Game 2001). Landings declined precipitously to 1,362 t in 1982, and the Bristol Bay fishery was closed in 1983 (Alaska Department of Fish and Game 2001) and more recently in 1994. Overall, declines in stock are considered to be a function of both anthropogenic and environmental effects (Loher et al. 1998). Specific reasons for population declines include an assortment of factors such as overfishing due to serial depletion (Orensanz et al. 1998), brood mortality due to parasites (Kuris et al. 1991), viral and microsporidian infections (Sparks and Morado 1985), climate change combined with overharvesting (Finney et al. 2000), temperature dependent growth, overharvesting based on incorrect population estimates (Stevens 1990), and changes in predator abundance (Anderson and Piatt 1999). Fisheries are now open as populations have stabilized (Orensanz et al. 1998). However, rebuilding strategies have had little effect on stock recovery (Loher et al. 1998). Prompted by the collapse of the P. camtschaticus (red king crab) fishery, fishermen started to target L. aequispinus (golden king crab) in the Aleutian Islands during the early 1980s and P. platypus (blue king crab). Paralithodes platypus occurs in a number of isolated pock- ets in Alaska, specifically Prince William Sound, the Kodiak region, Pribilof Islands, and St. Matthew Island (Orensanz et al. 1998). Currently, P. camtschaticus, P. platypus, and L. aequispinus remain among the most conservatively managed commercial fisheries in the world (T. Shirley, University of Alaska Fairbanks, Juneau, pers. comm.). These fisheries are regulated under the American Fisheries Act (Kruse et al. 2000). Regulations include a pre-specified harvest cap based on estimates of the effective spawning biomass of lithodids, a newly instated (2000) observer program, immediate catch size updates, pot limits, and enforcement vessels. For example, in 2000, fishing in Bristol Bay was opened on October 16 at 4:00 pm and closed by emergency order when quotas were met at 9:00 pm on October 20 (Alaska Department of Fish and Game 2001). Studies analyzing alternative rebuilding strategies are constantly being published and updated, taking into account important behavioral and biological information (e.g., Zheng et al. 1997a,b,c; Kruse et al. 2000). Table 8 contains cursory fishery comparisons, but up-to-date fishery information can be down- loaded from the internet (e.g., Alaska Department of Fish and Game 2001). In southern South America (Chile and Argentina) two sympatric species, Paralomis granulosa (the false centolla) and L. santolla (centolla), have constituted a mixed fishery since the 1930s (Lovrich 1996, Vinuesa et al. 1996, Lovrich and Vinuesa 1999). Originally L. santolla was the primary fishery with landings peaking in 1974 at 320 t. However, after 1984 landings of L. santolla precipitously declined (260 t), and the fishery for P. granulosa began to develop. Landings of P. granulosa peaked in 1996 (360 Crabs in Cold Water Regions: Biology, Management, and Economics 761 t), with L. santolla maintaining only bycatch status (1.5 t in 1996; for summary statistics and refer to Fig. 1 of Lovrich and Vinuesa 1999). The con- tinual violation of Argentine and Chilean fishery regulations is considered the main reason for the collapse. Transgressions, such as harvesting females and sublegal males, are frequent as effective controls are lacking (Vinuesa et al. 1995). The present reduction of L. santolla and P. granulosa landings, along with more restrictive regulations such as a shorter fishing season, has motivated the exploration and potential openings of new fisheries (G. Lovrich, pers. comm.). These include other lithodines such as L. confundens on the Atlantic coast of southern Argentina (49-53ºS), and P. spinosissima and P. f o rmosa in South Georgia Islands (Otto and Macintosh 1996).

Conclusion This paper provides an overview of what is known about a diverse and speciose (approximately 105 species) family of decapods. The family Lithodidae is divided into subfamilies Hapalogastrinae and Lithodinae. Members of the subfamily Hapalogastrinae are small, soft-abdomen bearing, solitary, intertidal inhabitants found only in the North Pacific Ocean. Members of the subfamily Lithodinae, in comparison, are large, fully cal- cified, social, deepwater inhabitants that have a pan-global distribution, also concentrated in the North Pacific Ocean (Tables 1 and 2). Although there is no single key differentiating all lithodids, good keys emphasizing the differences between lithodid genera have been published (Table 3). Evolutionary relationships are just beginning to be understood at the generic level (Fig. 1), however species-level relationships are poorly understood, especially in some species of Paralomis, Neolithodes, and Lithodes, where often only the type specimen is known (Table 4). Little is known about life history traits, including age of first reproduction of nonharvested species (Table 5). Only a cursory view of lithodid predators and prey is currently available (Table 6). Lithodids are host to several parasites such as the liparid fish Careproctus spp., and the rhizocephalan Briarosaccus callosus, as well as microsporidians and nemertean brood parasites which may be a major mortality source (Table 7) of harvestable species (Table 8, Fig. 2). The family Lithodidae is a large and diverse group of poorly understood animals. They possess complex life history patterns and live in en- vironments that tend to be inhospitable and inaccessible to biologists. Behaviors such as podding (Powell and Nickerson 1965a, Dew 1990), female preference, sperm limitation (Powell and Nickerson 1965b, Powell et al. 1973a, Sapelkin and Fedoseev 1986, Paul and Paul 1997, Paul and Paul 2001a) and slow growth rates (Paul 1992, Lovrich 1997) are important keys for fisheries management tools, as well as biodiversity estimates. Many species with high morphological similarity are known only from holotypes or a single sex, thus named species may represent natural 762 Zaklan — Review of Lithodidae intraspecific variation or may be indicative of the taxonomic infancy of this family, suggesting that there is a serious underestimation of oceanic biological diversity (Miya and Nishida 1997, Etter et al. 1999). However, due to their abyssal life-styles, discerning valid species from those vari- ants that are merely natural morphological extremes is often a difficult task. In this age of molecular systematics and increased taxonomic interest, future genetic analysis may aid in the basic goals of understanding species numbers, diversity, distributions, and evolutionary history and trajectories of lithodids.

Acknowledgments Editing was kindly provided by Drs. Felix Sperling, Brian Chatterton, John Spence, Tom Shirley, and Jody Martin. This paper benefited greatly from a review by Dr. Tom Shirley and authoritative help from Drs. Elliot Dawson, Enrique Macpherson, and Gustavo Lovrich. I would like to thank John Duff for creating the family Lithodidae internet site, Krystal Rypien for laboratory assistance, and the interlibrary loan department at the University of Alberta for extensive literature searches. Funding for this project was provided by the M&D philanthropic foundation of George and Evelyn Zaklan supplemented by NSERC grants to Drs. Felix Sperling and A.R. Palmer (P7245). Crabs in Cold Water Regions: Biology, Management, and Economics 763

Figure 1. Most parsimonious tree based on combined analysis of partial sequences from four mitochondrial data sets (12S, 16S COI, COII) and one nuclear data set (28S). Numbers at nodes represent bootstrap values for each of three analyses. First, maximum parsimony, a method which assumes the least changes over time. Second, maximum likelihood based on general times reversal model and site rate analysis, a method that accommodates rate heterogeneity across sites of each gene. Third, maximum likelihood invariant/gamma that accommodates invariant sites. Letters at nodes represent the following: FL = family Lithodidae, SH = subfamily Hapalogastrinae, SL = subfamily Lithodinae, FP = family Paguridae, FD = family Diogenidae, SP = superfamily Paguroidea. Outgroup taxa are Emerita analoga (superfamily Hippoidea) and Clibinarius vittatus (superfamily Paguroidea, family Diogenidae). 764 Zaklan — Review of Lithodidae

Figure 2. Harvest size, in millions of pounds, of king crabs (Paralithodes camtschaticus, P. platypus, and Lithodes aequispinus) landed in Alaska (Bristol Bay, Pribilof Islands, St. Matthew Island, Aleutian Islands, southeastern Alas- ka, and Norton Sound) between the years 1970 and 2000 (Alaska Depart- ment of Fish and Game 2001). Crabs in Cold Water Regions: Biology, Management, and Economics 765

Table 1. Location and distribution summaries of the family Lithodidae (update of Dawson 1989).

Location (Pacific, Atlantic, Taxon Authority Indian, Antarctic, Arctic oceans) Depth (m)a

Subfamily Ortmann Lithodinae 1901

Acantholithus Haan West Pacific: Japan, New Zealand Shallow and deep hystrix 1849 (Dawson 1989): species uncertain, water (Dawson 1989) see Tables 2-4 for details

Cryptolithodes Miers Northwest Pacific: Japan (Miers 50-60 (Kim and Hong expansus 1879 1879), Minyako, Rikuzen 2000) Province, Aomori near Hanaguri Cape between Aomori and Gutago Cape and between Benten Island and Cape Kurosaki (Makarov 1962); Korean coast on rocky bottoms in the sublittoral region (Kim and Hong 2000)

C. sitchensis Brandt Northeast Pacific: Sitka, Intertidal to 37 1853 Alaska (Makarov 1962) south (U.S.N.M. collection) to San Diego, California, U.S.A. (Bowman 1972)

C. typicus Brandt Northeast Pacific: Monterey, Low intertidal to 45 1848 California (Schmitt 1921); to (Hart 1965) Amchitka Island, Alaska, U.S.A. on rock-rubble bottom (Barr 1973)

Glyptolithodes (Faxon East Pacific: (07º9¢N, 80º50¢W), 245-580 (Bahamonde cristatipes 1893) Iquique, Chile (25º11¢S,70º31¢W; 1967), 245-800 (Haig Bahamonde 1967); south of Banco 1974) de Mancora, Peru (del Solar 1972), 03º51¢S, 81º18¢W (del Solar 1981); to Puerto Chicama, Mexico (07º42¢S, 80º26¢W), mud or rocky bottoms (Haig 1974); northern range of south California (Zaklan 2001)

Lithodes Benedict North Pacific: Bering Sea, Pribilof 315-730 (Makarov aequispinus 1895 Islands (Benedict 1895); Sea of 1962), 77-366 (Butler Okhotsk, Japan, 1895); east of and Hart 1962), 400- Siwoya Cape (Makarov 1962); to 900 (Hiramoto and south B.C., Canada, in the upper Sato 1970), 500-600 continental slope (Butler and (Sakai 1976), up to 742 Hart 1962); west Sagami Bay (U.S.N.M. collection) (Hiramoto and Sato 1970); off Shioya-zaki, off Matsushima, off Enoshima (Sakai 1976); and Suruga Bay, Japan (Suzuki and Sawada 1978)

L. couesi Benedict North Pacific: Bering Sea, 542-1,125 (Makarov 1895 north of Unalaska near the 1962), 695-820 Shumagin Islands, Alaska (Benedict (Takeda 1974), 258- 1895); to San Diego, California 1,829 (Hart 1982) (Makarov 1962); N.W. far off Midway Island (32º03.8¢N, 172º50.2¢E); Kushiro, Shioya-zaki (Takeda 1974); Hokkaido and off Onahama, Japan (Sakai 1976) 766 Zaklan — Review of Lithodidae

Table 1. (Continued.) Location and distribution of Lithodidae.

Location (Pacific, Atlantic, Taxon Authority Indian, Antarctic, Arctic oceans) Depth (m)a

L. confundens Macpherson Southwest Atlantic: south of 50-119 (Macpherson 1988c Falkland Islands (54º02¢S, 1988c) 58º40¢W), to Strait of Magellan (Punta Arenas), muddy bottoms (Macpherson 1988c)

L. ferox Filhol 1885 Atlantic: coasts of Mauritania 160 and 1,013 and Namibia, continental slope (Macpherson 1988c), off west coast of Africa continental shelf (22º03¢N to 28º16¢S), and 300-350 (Abello and South American coast of Saint Macpherson 1991) Helena, Brazil (36º29.6¢S to 53º46.7¢W), on muddy bottoms (Macpherson 1988c)

L. longispina Sakai 1971 West Pacific: off Matsushima, 600 (Sakai 1971), 400- Miyagi Prefecture; off 900 (Hiramoto 1974) Kominato, Chiba Prefecture (Sakai 1971); off Midway Island (32º02.9¢N, 172º45.3¢E; Takeda 1974); off Sendai, Boso Peninsula and Sagami Bay, Japan (Hiramoto 1974); South Pacific, Guam (Dawson 1989)

L. maja (Linnaeus North Atlantic: Shetland Islands, 95-532 (Hansen 1908), 1758) Scotland; Faroe Islands, 40-500 (Makarov England; Belgium, Holland; 1962), 65-790 (Will- Norway; Murman Sea (as far as iams 1984), 4-200 Teriberka), northernmost locality (Macpherson 1988c) 74º25¢N, 17º36¢E, Iceland, Greenland; coast of North America from Newfoundland to 40ºN (Hansen 1908)

L. mamillifer Macpherson West Indian Ocean: Mozambique 550-800 (Macpherson 1988 Channel between Madagascar 1988a) and Africa (Kensley 1977; as L. murrayi); La Reunion, Madagascar (22º18.9 S-43º01.1¢E), and off Natal (28º00¢S-32º46¢E), South Africa, in mud (Macpherson 1988a)

L. manningi Macpherson Central Atlantic: Dominica, 640-777 (Macpherson 1988 French Guiana (Macpherson 1988c) 1988c)

L. murrayi Henderson Southwest Pacific, southeast Coast of Namibia 360- 1888 Atlantic, south Indian: south 800, Fondos de fango New Zealand (Yaldwyn and 120-810 (Macpherson Dawson 1970); Fondos de fango, 1983), 75-700 (Takeda Islas Posesion, Prince Edwards, and Hatanaka 1984), Macquarie and Crozet Islands of Indian Ocean 80-1,015 South Africa and off Namibia (Miquel et al. 1985), (18º11¢S and 28º16¢S), south of 35-200 (Macpherson Chile (Macpherson 1983) 1988c)

aU.S.N.M. = Information was obtained from the United States National Museum of Natural History collections. Crabs in Cold Water Regions: Biology, Management, and Economics 767

Table 1. (Continued.)

Location (Pacific, Atlantic, Taxon Authority Indian, Antarctic, Arctic oceans) Depth (m)a

L. nintokuae Sakai 1978 Northwest Pacific: north of 450-1,070 (Dawson and Nintoku Seamount, Emperor Yaldwyn 1985b) Seamount Chain, Japan (Sakai 1978); northwest of Midway/ Hawaiian Islands Ridge (21º23¢N, 158º14¢W; 32º03.8¢N, 172º50.2¢E; Dawson and Yaldwyn 1985b)

L. panamensis Faxon 1893 East Pacific: Colombia 680-850 (del Solar 1972) (07º31¢N, 79º14¢W; Faxon 1895); to Peru (03º48¢S, 81º22¢W; 07º59¢S, 80º22¢W; 17º34¢S, 71º55¢W), on hard bottoms (del Solar 1972, Haig 1974); Gulf of Panama (Wicksten 1989); northern range Baja California, U.S.A. (Dawson 1989)

L. richeri Macpherson Southwest Pacific: New Caledonia,Trapped outside coral 1990 southeast Australia (Macpherson reef, depth unrecorded southeast Australia (Macpherson (Macpherson 1990) 1990)

L. santolla (Molina Southeast Pacific and southwest Intertidal to 700 (Boschi 1782) Atlantic: south of South America, et al. 1984), concen- Strait of Magellan, north (Takeda trate between 10 to 50 and Hatanaka 1984); to (Macpherson 1988c) Talcahuano, Chile (36º41¢S; Hernandez 1985); east coast of South America, Tierra del Fuego to Uruguay (34ºS, Vinuesa et al. 1996)

L. turkayi Macpherson Southeast Pacific and southwest 70 (Campodonico and 1988 Atlantic: Falkland Islands, U.K. Guzman 1972), to 581 and coast of Chile from Tierra (Revuelta and Andrade del Fuego north to 31º56¢S, 1978) 71º38¢W (Campodonico and Guzman 1972 (as L. murrayi), Revuelta and Andrade 1978, Macpherson 1988c)

L. turritus Ortmann Northwest Pacific: Sagami Bay, 200 (Sakai 1971), to 812 1892 off Boso Peninsula, Chiba (Macpherson 1990) Prefecture to Tosa Bay, Japan (Sakai 1971); Philippines on soft sandy-mud bottoms (Macpherson 1990); east China Sea, Taiwan (Wu et al. 1998)

L. unicornis Macpherson Southeast Atlantic: off southwest 934-936 (Macpherson 1984 Africa, Valdivia Bank (24º43.7¢S, ) 1984) 06º24.3¢E), on muddy bottoms (Macpherson 1984

L. wiracocha Haig 1974 Southeast Pacific: 03º48¢S, 620-800 (del Solar 1972, 81º22¢W; 07º59¢S, 80º22¢W ) Haig 1974) (del Solar 1972); SW Banco de Mancora Peru, mud bottom (Haig 1974) 768 Zaklan — Review of Lithodidae

Table 1. (Continued.) Location and distribution of Lithodidae.

Location (Pacific, Atlantic, Taxon Authority Indian, Antarctic, Arctic oceans) Depth (m)a

Lopholithodes (Faxon East Pacific: 07º31¢30¢ N, 79º14¢W; 680-935 (Faxon 1893, diomedeae 1893) 07º21¢N, 79º35¢W (Faxon 1895); 1895; del Solar 1972) hard mud bottoms, Gulf of Panama to Peru (03º48¢S, 81º22¢W; 10º01¢S, 79º10¢W; del Solar 1972)

L. foraminatus (Stimpson Northeast Pacific: San Diego, Intertidal to 547 (Hart 1859) California to Banks Island, Hecate 1982, U.S.N.M. strait, B.C., Canada (53º40¢N, collection) 130º30¢W); on muddy bottoms (Hart 1982); north to Aleutian Islands and Bering Sea (Dawson 1989, Wicksten 1989)

L. mandtii Brandt 1848 Northeast Pacific: Sitka, Intertidal to 137 (Hart Alaska to Monterey California, 1982) U.S.A. (Makarov 1962); rocky habitats with strong currents, juveniles found under rocks during extremely low tides (Hart 1982, Jensen 1995)

L. odawarai Sakai 1980 Northwest Pacific: Sagami Bay, 240-280 (Sakai 1980) Japan (Sakai 1980)

Neolithodes (Smith 1882) Northwest Atlantic: east coast 650-1,900 (Macpherson agassizii of North America between 1988c) 34º39¢N, 75º14¢W (Smith 1882); and 07º22¢N, muddy bottoms (Macpherson 1988c); Gulf of Mexico and off Suriname and French Guiana (Dawson 1989)

N. asperrimus Barnard 1946 Southeast Atlantic: off Saldanha 870 and 1,007 (Barnard Bay and Cape Point, southern 1946), 530 and 615 South Africa (Barnard 1946); (Macpherson 1983), area del Cabo, Namibian coast 600-2,000 (Macpher- (18º11¢S to 28º16¢S; Macpherson son 1988c) 1983); 13º46 S, 47º33 E; 13º48 S, 47º29¢E (Macpherson 1988a); Ivory coast, West Africa on muddy bottoms (Macpherson 1988c)

N. brodiei Dawson and Southwest Pacific: New Zealand 832 (Dawson and Yaldwyn and southeast Australia, Yaldwyn 1970) 1970 Campbell plateau (50º58¢S, 173º57¢E; Dawson and Yaldwyn 1970)

N. capensis Stebbing Southeast Atlantic, Indian Ocean: 1,570-2,745 (Kensley 1905 south South Africa, off Cape 1968), 1,480-3,200 Point (Kensley 1968); on muddy (Macpherson 1988c) bottoms (Macpherson 1988c)

aU.S.N.M. = Information was obtained from the United States National Museum of Natural History collections. Crabs in Cold Water Regions: Biology, Management, and Economics 769

Table 1. (Continued.)

Location (Pacific, Atlantic, Taxon Authority Indian, Antarctic, Arctic oceans) Depth (m)a

N. diomedeae Benedict East Pacific: south Chile (42º36¢S 1,382-2,454 (Haig 1895 and 45º35¢S; Benedict 1895); 1974), 1,100-2,000 Scotia Sea, Antarctica, north (Baez et al. 1986) to Mexico and southern California, U.S.A. (Dawson 1989); Gulf of Panama (Wicksten 1989)

N. grimaldii (A. Milne North Atlantic: Iceland, off east 1,065 (Hansen 1908), Edwards coast of Canada and U.S.A. 1,267-3,000 (Macpher- and (Hansen 1908); north of 35º23¢N, son 1988c), to 3,207 Bouvier Greenland and western Ireland, (U.S.N.M. collection) 1894) south to Bay of Biscay, Canary Islands, and Cape Verde, Spain on muddy bottoms (Macpherson 1988c)

N. martii Birstein and Southwest Atlantic: near South 305-650 (Birstein and Vinogradov America (53-54ºS, 34-36ºW; Vinogradov 1972 1972 Birstein and Vinogradov 1972)

N. nipponensis Sakai 1971 Northwest Pacific: Mikawa 200-600 (Sakai 1971, Bay and Kii Peninsula, Japan 1976) (Sakai 1971,1976)

N. sp. nov. Dawson Indian Ocean: Bay of Bengal Deep water (Dawson and and Madagascar (Dawson 1989) 1989) Yaldwyn unpubl. manuscr.

N. vinogradovi Macpherson Southwest Pacific and southeast 1,600 (Macpherson 1988 Indian Ocean: 31º50¢43¢¢S, 1988c), to 2,100 87º22¢27¢¢E (Macpherson 1988c); (Macpherson 1990) New Caledonia, Coral Sea (Macpherson 1990)

Paralithodes (A. Milne North Pacific: Sea of Japan, south Intertidal to 66 brevipes Edwards to Cape Povorotnyi, Sea of Okhotsk, (Makarov 1962, and east Kamchatka, Russia; south U.S.N.M. collection) Lucas 1841) Bering Sea, Aleutian Islands, U.S.A. (Makarov 1962)

P. californiensis (Benedict Northeast Pacific: Monterey Bay to 148-306 (Schmitt 1921), 1895) San Diego, California (Schmitt to 349 (U.S.N.M. 1921, Anderson and Cailliet 1974) collection)

P. camtschaticus (Tilesius North Pacific: Bristol Bay, Alaska, 3-366 small juveniles 1815) U.S.A. (Benedict 1895); Bering Sea found intertidally to Sea of Japan (Marukawa 1930); among rocks and algae Hokkaido, Japan; Cape Gamova, (Marukawa 1930, Sea of Okhotsk, eastern Kamchatka Jensen 1995) to Cape Olyutorsk, Russia; Aleutian Islands and Norton Sound, U.S.A. to Queen Charlotte Islands, B.C., Canada (Makarov 1962); Korea (Kim 1970); sand or mud bottoms (Jensen 1995) age 0-1 located on complex bottoms (Loher and Armstrong 2000) 770 Zaklan — Review of Lithodidae

Table 1. (Continued.) Location and distribution of Lithodidae.

Location (Pacific, Atlantic, Taxon Authority Indian, Antarctic, Arctic oceans) Depth (m)a

P. platypus (Brandt North Pacific: Sea of Japan; south to 12-500 (Makarov 1962) 1850) Cape Gamova, Sea of Okhotsk, east Kamchatka, Russia; Bering Strait (Makarov 1962); Alaska; Tartary Bay, Vladivostok; Sakhalin, Kurile Island, Kitami, Japan; Korea (Sakai 1976)

P. rathbuni (Benedict Northeast Pacific: San Simeon Bay, 367-402 (Schmitt 1921), 1895) California, U.S.A. (Benedict 1895); 201 (Wicksten 1987) to Baja California, Mexico (Wicksten 1987)

Paralomis Henderson Southeast Atlantic, Indian Ocean: 560 (Takeda 1974) aculeata 1888 Prince Edward (Takeda 1974); and Crozet Islands, south of South Africa (Dawson 1989)

P. africana Macpherson Southeast Atlantic: Fondos de fango, 570-770 (Macpherson 1982 Namibian coast (20º31¢S to 24º42¢S), 1988c) mud habitat (Macpherson 1982, 1983)

P. anamerae Macpherson Southwest Atlantic: north of Falkland 132-135 (Macpherson 1988 Islands (Malvinas), off Argentina 1988c) (Macpherson 1988c)

P. aspera Faxon 1893 East Pacific: 07º06’N, 80º34¢W 560-1,271 (Faxon 1893, (Faxon 1895); Panama to Peru 1895; del Solar 1972) (03º48¢S, 81º20¢W; del Solar 1972, Haig 1974)

P. birsteini Macpherson Antarctic Ocean: 67º29¢S, 79º55¢W (Macpherson 1988b) 1988 500-1,080 (Macpher- son 1988b)

P. bouvieri Hansen Northeast Atlantic: between 1,345-1,454 (Hansen 1908 Greenland and Iceland (65º24¢N, 1908), 1,460-4,152 29º00¢W; Hansen 1908); northeast (Macpherson 1988c), U.S.A. and off SW Ireland 889-1,500 (Pohle (Macpherson 1988c); 1992b) Canadian Atlantic(43ºN, 59ºW; Pohle 1992b)

P. ceres Macpherson Indian: Arabian Sea (22º22¢12¢¢N, 1,189-1,354 (Macpher- 1989 59º57¢30¢¢E; Macpherson 1989) son 1989)

P. cristata Takeda and Northwest Pacific: Suruga Bay off 750 (Takeda and Ohta Ohta 1979 Osaki (Takeda and Ohta 1979); 1979), to 1,100 (Sakai abyssal valley off Gamoda-misaki, 1987) Japan (Sakai 1987)

P. chilensis Andrade Southeast Pacific: central Chile to 400-420 (Andrade 1980) 1980 Peru (31º56¢S to 29º50¢S; Andrade 1980)

P. cristulata Macpherson Southeast Atlantic: west Africa, off 261-650 (Macpherson 1988 Guinea Bissau and Senegal, on 1988c) muddy bottoms (Macpherson 1988c)

aU.S.N.M. = Information was obtained from the United States National Museum of Natural History collections. Crabs in Cold Water Regions: Biology, Management, and Economics 771

Table 1. (Continued.)

Location (Pacific, Atlantic, Taxon Authority Indian, Antarctic, Arctic oceans) Depth (m)a

P. cubensis Chace 1939 Central west Atlantic, West Indies: 439-550 (Chace 1939), east of Havana, Cuba (Chace 1939); 329-730 (Macpherson east of Florida, Gulf of Mexico and 1988c) northern Brazil (Macpherson 1988c)

P. debodeorum Feldmann Southwest Pacific: middle to late 200-300 (Feldmann 1998 Miocene fossils (10 mya), of 1998) Motunau Beach, North Canterbury, New Zealand (Feldmann 1998)

P. dofleini Balss 1911 Northeast Pacific: off Sendai Bay 470-780 (Takeda 1974) off Kominato, Sagami Bay, Japan (Takeda 1974, Sakai 1976)

P. erinacea Macpherson East Atlantic: west Africa, off Guinea 251-900 (Macpherson 1988 Bissau and Ivory Coast on muddy 1988c) bottoms (Macpherson 1988c)

P. formosa Henderson Southwest Atlantic: off Rio Plata 400-1,599 (Macpherson 1888 (Takeda 1974); off Argentina and 1988c) Uruguay, South Georgia and South Orkney Islands (Macpherson 1988c)

P. granulosa (Jacquinot Southwest Atlantic and southeast Intertidal to 100 1852) Pacific: southern South America (Takeda and Hatanaka from Rio de Janeiro, Brazil through 1984) Falkland Islands, U.K. and Magellanic district, Argentina to the vicinity of Chiloe Island, Chile (Takeda and Hatanaka 1984); juveniles found in kelp beds (Hoggarth 1993)

P. grossmani Macpherson Central west Atlantic: coast of 770 (Macpherson 1988c) 1988 French Guiana (Macpherson 1988c)

P. haigae Eldredge Central west Pacific: Adelup Point, 400-730 (Eldredge 1976) 1976 Guam (Eldredge 1976); New Caledonia, Samoa Islands (Macpherson 1990)

P. heterotuber- Yumao Northwest Pacific: east China Sea 860-890 (Yumao et al. culata et al. (30º26¢N, 128º53¢E; Yumao et al. 1984) 1984 1984)

P. hystrixoides Sakai 1980 Northwest Pacific: Sagami Bay off 750-1,100 (Sakai 1987) Daiozaki, Mie Prefecture (Sakai 1980); Abyssal valley off Gamoda- misaki, Tokushima, Japan (Sakai 1987)

P. hystrix Haan 1846 Northwest Pacific: Tokyo to Tosa 300-600 (Takeda 1974), Bay (Takeda 1974); Boso Peninsula, 230-300 (Sakai 1980) Chiba Prefecture, south to Nagasaki, Japan (Sakai 1976)

P. inca Haig 1974 Southeast Pacific: Peru (06º31ºS, 620-800 (Haig 1974) 81º01¢W); mud and sand (Haig 1974) 772 Zaklan — Review of Lithodidae

Table 1. (Continued.) Location and distribution of Lithodidae.

Location (Pacific, Atlantic, Taxon Authority Indian, Antarctic, Arctic oceans) Depth (m)a

P. indica Alcock and Indian: Travancore coast, India 786 (Alcock and Anderson (Alcock and Anderson 1899) Anderson 1899) 1899

P. investigatoris Alcock and Indian: Travancore coast, India 786 (Alcock and Anderson (Alcock and Anderson 1899) Anderson 1899) 1899

P. japonica Balss 1911 Northwest Pacific: Sagami Bay, Shallow water (Dawson Japan (Balss 1911, Sakai 1971) 1989)

P. jamsteci Takeda and Northwest Pacific: Minami-Ensei 710 (Takeda and Hashimoto Knoll (28º23¢N, 127º38¢E), in the Hashimoto 1990) 1990 mid-Okinawa trough, Japan (Takeda and Hashimoto 1990)

P. kyushup- Takeda Northwest Pacific: northern part of 340-460 (Takeda 1985) alauensis 1985 the Kyushu-Palau submarine ridge, Japan (26º47¢N, 135º20¢E to 26º48¢N, 135º21¢E; Takeda 1985)

P. longidactyla Birstein and Southwest Atlantic: near mouth of 485-500 (Macpherson Vinogradov River Plate (35º34¢S, 52º40¢W), 1988c) 1972 Uruguay (Birstein and Vinogradov 1972)

P. longipes Faxon 1893 East Pacific: 05º26¢N, 86º55¢W 760-1,409 (Faxon 1893, (Faxon 1895); Peru (07º59¢S, 1895; del Solar 1972) 80º22¢W; 16º29¢S, 73º33¢W; del Solar 1972); off San Diego, California, U.S.A., hard bottom (Haig 1974)

P. manningi Williams Northeast Pacific: San Clemete Basin, 1922 (Williams et al. et al. 2000 southern California (Williams et al. 2000) 2000)

P. medipacifica Takeda 1974 Northwest Pacific: off Midway, 695-820 (Takeda 1974) Japan (Takeda 1974)

P. microps Filhol 1884 Northeast Atlantic: Bay of Biscay, 1,480 (Macpherson Japan (Takeda 1974) 1988c)

P. multispina (Benedict North Pacific: Queen Charlotte 1,603 (Benedict 1895) Islands, B.C., Canada (Benedict 1895),1,143-1,603 1895); Shumagin Islands, Alaska (Rathbun 1904), 1,125- to San Diego, California, U.S.A.; 1,577 (Makarov 1962), west Bering Sea, Kamchatka, Russia 600-830 (Sakai (Makarov 1962); off Hokkaido, 1971),830-1,665 (Hart Miyagi Prefecture, Chiba Prefecture, 1982) off Manazuru and Enoshima, Sagami Bay, Japan (Sakai 1971)

P. ochthodes Macpherson Indo-Pacific: Sulawesi (Celebes), 1,281 (Macpherson 1988 Islands, Indonesia (04º43¢S, 1988b) 121º23¢E; Macpherson 1988b) aU.S.N.M. = Information was obtained from the United States National Museum of Natural History collections. Crabs in Cold Water Regions: Biology, Management, and Economics 773

Table 1. (Continued.)

Location (Pacific, Atlantic, Taxon Authority Indian, Antarctic, Arctic oceans) Depth (m)a

P. otsuae Wilson 1990 Southeast Pacific: off Mejillones 1,740 (Wilson 1990) del Sol, Chile (22º55¢S, 70º46¢W; Wilson 1990)

P. pacifica Sakai 1978 Northwest Pacific: north of Nintoku 800 (Sakai 1978) Seamount, Japan (42º20¢N, 170º50¢E; Sakai 1978)

P. papillata (Benedict East Pacific: southern California 712-744 (Haig 1974) 1895) (Benedict 1895); to Peru (06º31¢S, 81º01¢W), in mud and sand (Haig 1974)

P. pectinata Macpherson West central Atlantic: Caribbean 1,409-1,629 (McPherson 1988 Sea, off Isla Margarita, Venezuela 1988c) (Macpherson 1988c)

P. phrixa Macpherson Southeast Pacific: coast of Peru ) 1,815-1,860 (Macpher- 1992 (04º10’S, 81º27’W; Macpherson son 1992) 1992

P. roeleveldae Kensley 1981 Southwest Indian Ocean: east 625-900 (Kensley 1981) coast of South Africa (30º32’S, 30º52’E; Kensley 1981)

P. seagranti Eldredge Southwest Pacific: Double Reef area 250-620 (Eldredge 1976 and Tanguisson Point, northwest 1976)750 (Macpherson coast of Guam (Eldredge 1976) 1990)

P. serrata Macpherson West central Atlantic: Caribbean sea 1,100 (Macpherson 1988 off Colombia (Macpherson 1988c) 1988c)

P. shinkaimaruae Takeda 1984 Southwest Atlantic: Bromley Plateau 668 (Takeda and (31º13¢S, 34º49¢W; Takeda and Hatanaka 1984) Hatanaka 1984)

P. sp. nov. cf. Macpher- Southwest Pacific: Louisville Ridge, 731-1,097 (Webber, son 1990 east of the North Island, New pers. comm.) fide Zealand (Webber, pers. comm.) Webber, and Dawson unpubl. manuscr.

P. sp. nov. Macpherson Southwest Pacific: New Caledonia Depth unrecorded 1990 (Macpherson 1990) (Macpherson 1990)

P. spectabilis Hansen Northwest Atlantic and subantarctic: 1,345-1,786 (Hansen 1908 Scott Island in Ross Sea, near 1908), 1,470-2,075 Iceland and Greenland (64º44¢N, (Macpherson 1988c) 32º32¢W; Hansen 1908)

P. spinosissima Birstein and Southwest Atlantic, Antarctic Ocean: 215-650 (Birstein and Vinogradov South Georgia Island (53-54ºS, Vinogradov 1972), 1972 34-36ºW; Birstein and Vinogradov 132-650 (Macpherson 1972); between Burwood Bank and 1988c) Falkland Islands (Macpherson 1988c) 774 Zaklan — Review of Lithodidae

Table 1. (Continued.) Location and distribution of Lithodidae.

Location (Pacific, Atlantic, Taxon Authority Indian, Antarctic, Arctic oceans) Depth (m)a

P. stella Macpherson West Indian Ocean: Madagascar and 350-750 (Macpherson 1988 La Reunion (19º41¢S, 54º08¢E; 1988a) Macpherson 1988a)

P. truncatispinosa Takeda and Northwest Pacific: continental slope 642-840 (Takeda and Miyake of East China Sea, stone and sandy Miyake 1980) 1980 mud (Takeda and Miyake 1980)

P. tuberipes Macpherson Southeast Pacific: off Huichas Depth unrecorded 1988 Islands, Puerto Aguirre, Chile (Macpherson 1988b) (45º10¢S, 73º33¢W; Macpherson 1988b)

P. verrilli (Benedict North Pacific: Bering Sea, Pribilof 1,238-1,480 (Makarov 1895) Islands (Benedict 1895); south to 1962), 1,238-2,379 Cortez Bank, California, U.S.A. (Hart 1982), 850-1,250 (Makarov 1962); coast of Nemuro (Sakai 1987) Hokkaido (Sakai 1976); Kumano- nada, off Daiozaki, Mie prefecture, Abyssal valley off Gamoda-misaki, Japan (Sakai 1987)

P. zealandica Dawson and Southwest Pacific: Chatham Rise, 640 (Dawson and Yaldwyn New Zealand (44º18¢S, 174º31¢E), Yaldwyn 1971) 1971 fine sandy mud (Dawson and Yaldwyn 1971)

Phyllolithodes Brandt 1848 Northeast Pacific: Unalaska, Alaska Subtidal to 183 (Hart papillosus to Monterey, California, U.S.A. 1982), some juveniles (Makarov 1962); rocky areas with low intertidal (Jensen currents (Jensen 1995) 1995)

Rhinolithodes Brandt 1848 Northeast Pacific: Kodiak Island, 6-73 (Hart 1982), up to wosnessenskii Alaska to Crescent City, California, 102 (Chapter 2) generally on rocky bottoms and on shells (Makarov 1962)

Sculptolithodes Makarov Northwest Pacific: Sea of Japan, 20-35 (Makarov 1962) derjugini 1934 near Silant’ev Bay and in the area of Nel’ma Bay, Andreev Bay (Ussuri Bay), and Rishiri Island, rocky bottoms (Makarov 1934, 1962); Hokkaido, Japan (Sakai 1976)

Subfamily Ortmann Hapalo- 1901 gastrinae

Acantholithodes (Stimpson Northeast Pacific: off Moorovskoy Intertidal to 245 hispidus 1860) Bay, Alaska to Monterey, (Benedict in litt. California (Hart 1982) U.S.N.M. collection)

aU.S.N.M. = Information was obtained from the United States National Museum of Natural History collections. Crabs in Cold Water Regions: Biology, Management, and Economics 775

Table 1. (Continued.)

Location (Pacific, Atlantic, Taxon Authority Indian, Antarctic, Arctic oceans) Depth (m)a

Dermaturus Brandt 1850 North Pacific: Bering Sea to the Low intertidal to 72 mandtii Pribilof Islands, Alaska, U.S.A. (Makarov 1962) along the Asiatic coasts north to Cape Olyutorsk, eastern shores of Kamchatka; Sea of Okhotsk, north Sea of Japan in rocky areas (Makarov 1962); among kelp holdfasts and shell rubble in cavities (Jensen 1995)

Hapalogaster Stimpson Northeast Pacific: Washington, Intertidal to 15 cavicauda 1859 California, U.S.A., and Mexico (Dawson 1989) (Dawson 1989); clings tightly to the undersides of rocks (Jensen 1995)

H. dentata (Haan 1849) Northwest Pacific: south Japan, Intertidal to 180 north to Aomori and Hakodate, (Makarov 1962) Japan, Sea of Japan, north to Peter the Great Bay, Russia (Makarov 1962); Korean coasts (Kim 1970); intertidal and subtidal cobble rocky shores, in groups clinging to or hiding under cobbles (Goshima et al. 1995)

H. grebnitzkii Schalfeew North Pacific: eastern shores Intertidal to 90 1892 of Kamchatka, Sea of Okhotsk, (Makarov 1962) Russian shores of Sea of Japan, to Sibiryakov Island; Bering Sea, north to Bering Strait, west coast of North America from Aleutians, Alaska south to Humboldt Bay, California, U.S.A., on rocky bottoms (Makarov 1962)

H. mertensii Brandt Northeast Pacific: Atka, Alaska to Low intertidal to 55 1850 Puget Sound, Washington, U.S.A. (Hart 1982) (Makarov 1962); found between algae-covered rocks (Jensen 1995)

Oedignathus (Stimpson North Pacific: Japan, from the Middle intertidal to 15 inermis 1860) Tsushima Strait to Aomori (Tsugaru (Hart 1982) Strait); Patrocles Bay, Peter the Great Bay, Russia, west to Unalaska, Alaska south to Pacific Grove, California, U.S.A. (Makarov 1962); Korea (Kim 1970)

Placetron Schalfeew Northeast Pacific: Aleutian Islands, Intertidal to 110 wosnessenskii 1892 Alaska (Makarov 1962); south to (Hart 1982) Puget Sound, Washington, U.S.A. among anemones (Metridium; Hart 1982); on vertical rock faces and among boulders (Jensen 1995)

P. forcipatus (Benedict Northeast Pacific: Parry Passage, Shallow water 1895) Graham Island, B.C., Canada (Dawson 1989) (Benedict 1895): species uncertain, see Tables 2-4 for details 776 Zaklan — Review of Lithodidae

Table 2. A summary of global distribution of lithodid species.

North/south/ East/ Number % of a,b a,b Ocean central west of species species

Pacific 71 (62/9) 68 (65/100) North 51 (42/9) 49 (44/100) East 33 (25/8) 31 (26/89) West 28 (24/4) 27 (25/45) South 25 24 East 16 15 West 12 11 Central 1 1

Atlantic 29 28 North 7 7 East 4 4 West 3 3 South 17 16 East 8 8 West 11 10 Central 7 7

Indian 11 11

Antarctic 3 3

The subfamily Hapalogastrinae resides only in the North Pacific Ocean. aAs species may reside in multiple areas the numbers do not total 105 species and the percentage does not total 100%. Localities are obtained from Table 1. bParentheses denote percentage lithodid family members (subfamily Lithodinae/subfamily Hapalogas- trinae) that inhabit each of the specified oceans. Crabs in Cold Water Regions: Biology, Management, and Economics 777

Lovrich 1999 Hart 1971

Konishi and Shikatani 1998, 1999 MacDonald et al. 1957

Makarov 1962 Sandberg and McLaughlin 1998

Hart 1982 1994 Konishi and Taishaku

Macpherson 1988c Haynes 1984

Kozlov 1996

Neolithodes,

Reference

Lithodes

Paralomis granulosa

Neolithodes

Paralomis

, and

Lithodes

val characters of subfamily

)

only lithodid)

hystrix

Cryptolithodes

L. maja

) of Atlantic lithodids

, and

Paralithodes

Lithodes,

Comments

Paralomis

japonicus

Beagle Channel, Argentina and Chile

Morphological keys for zoeae and glaucothoe and chromatophore crabs in Japan

keys of

Lithodinae and one comparing zoeae of P and characters used to distinguish lithodid and pagurid zoeae and between stages I-IV of lithodid zoeae

of America

Paralomis

able 3. A list of taxonomic keys to members of the family Lithodidae.

axon Larvae or adult

Paguridae and

Anomura L Key for identification of Anomura and Brachyura zoeae in the Decapoda L Keys to the families of decapod crustacean larvae Lithodidae A Key to some lithodid species (

Anomura L Identification manual for larvae of commercially important Lithodidae key for zoeae (

Lithodidae A Keys to subfamilies and genera of lithodids as well as species level

Lithodidae A Keys to lithodid species of B.C., Canada Lithodinae L tables, one comparing lar Two

Lithodidae ALithodidae Keys to subfamilies, genera, and species ( L Key to species of lithodid zoeae of North Pacific Ocean;

Lithodidae A Keys to the species of lithodids found in the Pacific Northwest 778 Zaklan — Review of Lithodidae

illimas et al. 2000

Schalfeew 1892

Haynes 1982

Abe 1992 Dawson and Yaldwyn 1985a Dawson and Yaldwyn

Sakai 1971, 1976

Macpherson 1992

brevipes

zoeae Jensen et al. 1992

, and

larvae Campodonico and Guzman 1981

Reference

aequispinus

L. maja, L. santolla,

P. platypus P.

platypus

L. santolla

and

from

and

from its congeners

Paralomis

camtschaticus

granulosa

camtschaticus

L. aequispinus

manningi

tropicalis

species and based on rostral and carapace

, and

larvae

Lithodes

Hapalogaster

maja

Comments

Paralithodes

Key to genus

and

Features used for lithodid identification; key to lithodid genera; key to twelve characters they are divided into four groups: antarcticus (1988c) to distinguish

Differentiates between Differentiates

Differentiates between Differentiates Key to Japanese species of

Key to eastern Pacific species of

Modifications to key to species of Paralomis from Macpherson

A A

and L between Differentiates

and L Distinguishing larvae of

able 3. (Continued.) A list of taxonomic keys to members of the family Lithodidae.

axon Larvae or adult

Hapalogaster

Lithodes Paralithodes

Lithodes

Paralithodes

Paralomis Lithodes Paralithodes Paralomis

Paralomis

Paralomis Crabs in Cold Water Regions: Biology, Management, and Economics 779

cristata

Other species of Cryptolithodes

Paralomis (Takeda and Ohta (Takeda 1979)

species

size (mm) related

Maximum Closely male/female

Male: CL = 42, CW = 75 (Miyake 1982), female: CL = 25.84, R = 5.88, CW = 35.82 (Zaklan, pers. obs.)

Male: CL = 65, CW = 90, female: CL = 68, CW = 87 (Hart 1982)

Male: CL = 41, CW = 75, female: CL = 49, CW = 80 (Hart 1982)

Male: CL = 79.5, R = 6, CW = 98, female: CL = 89.5 (Haig 1974)

Paralomis

seudolithodes

(Sakai 1976)

comments

Considered hystrix

Includes P of Birstein and

Sato 1970)

Previous generic names and

Sakai 1971, 1976; Macpherson 1988c

map Distribution

Photo (Miyake 1982)

Drawings (Makarov 1962, Hart

Drawing (Makarov 1962, Hart 1982), photo (Schmitt

Drawing (Bahamonde 1967), dorsal, lateral and abdominal drawings (Haig 1974, Macpherson 1988c), photo (del Solar 1981)

or drawings

able 4. Photographs, distributions, maximum sizes, and general information concerning lithodids.

Scientific and Locations of vernacular photographs names Subfamily Lithodinae Acantholithus hystrix

Cryptolithodes expansus, Menko-gani

C. sitchensis, Sitka crab, 1982, Macpherson 1988c), turtle crab, photos (Schmitt 1921, umbrella crab Barr 1973, Jensen 1995)

C. typicus, butterfly crab, turtle crab 1921, Barr 1973, Jensen 1995)

Glyptolithodes cristatipes

Lithodes 780 Zaklan — Review of Lithodidae

L. aequispinus

Closely

L. santolla (Macpherson 1988c)

Lithodes turritus and (Hiramoto 1974)

from Dawson 1989.

species

size (mm) related

Maximum male/female

Male: CW = 220, female: CW = 192 (Hiramoto and Sato 1970)

CW = 85, female: CL = 105 (Lovrich and Vinuesa 1999) and Vinuesa

Male: CL = 105, CW = 103, female: CL = 115, CW = 113 (Takeda 1974) (Takeda

Male: CL = 145, R = 62, CW = 140 (Sakai 1987), female: CW = 125 (Macpherson 1988c) (Hiramoto 1974)

” as it Male: CL = 100

confundo

comments

Latin “ is thus confusion would arise (Mac- pherson 1988c)

Lithodes tropicalis A. Milne-Edwards 1883

Previous generic names and

Sakai 1971, 1976

Macpherson 1988c

Sakai 1971, 1976

Macpherson 1988c

Sakai 1971, 1976; Hiramoto 1974

map Distribution

Drawing (Makarov 1962,

(Sakai 1971, 1976,

Drawings (Macpherson 1988c)

Drawing (Makarov 1962, Hart 1982), photo (Schmitt

Sakai 1971, 1976, Somerton 1981)

Photo and drawings (Macpherson 1988c)

Drawing (Sakai 1971), photos (Takeda 1974; Sakai 1971, (Takeda 1976, 1987; Hiramoto 1974; (Sakai 1987) Macpherson 1990), drawing

or drawings

L. santollo

ferox

able 4. (Continued.) Photographs, distributions, maximum sizes, and general information concerning lithodids.

CL = carapace length (includes rostral length unless otherwise noted), CW = carapace width, R = rostrum length.

Macpherson 1988c maps are for Atlantic Ocean distributions only

For an exhaustive list see Dawson 1989. All uncited common Japanese names are obtained from Sakai 1976; other common names are

As a complete taxonomic account of each species is beyond the scope of this paper only selected junior synonyms are listed.

Scientific and Locations of vernacular photographs names L. aequispinus, Ibaragani Hart 1982), photo -modoki, golden king crab Miyake 1982)

L. confundens similar to

L. couesi, Kita-ibaragani, deep-sea or 1974, 1921, Takeda scarlet king crab

L. longispina Hari-ibaragani

a b c d Crabs in Cold Water Regions: Biology, Management, and Economics 781

and

(Macpher-

L. turkayi, L. unicornis L. ferox son 1988c)

L. murrayi (Faxon 1893)

species

size (mm) related

Maximum Closely male/female

aldwyn 1985b)

Male: CL = 110, Male: CL = 37, CW = 40, CW = 113, female: female: CL = 145,

(Macpherson 1988c)

female: CL = 106,

1988c)

Male: CL = 132.5, 1,500 g, female: CL = 93.5, 360 g (Miquel et al. 1985)

Male: CL = 116, CW = 123, R = 19

Male: CL = 190 (del Solar 1972), female: CL = 100,

. R.B. Male: CL = 94, CW = 103,

”

mamilla

comments

Female with reversed referring to the (Zaklan 2000) roundedcarapace’s CL = 97, CW = 95 CW = 158 (Macpherson protrusion 1988a) (Macpherson 1988a) Latin “ Named after Dr Manning (Macpherson 1988c) CW = 120 (Macpherson

abdominal asymmetry (Dawson and Yaldwyn (Sakai 1978), female: 1985b) CL = 115 (Dawson and

Male CL = 970 between tips of extended pereiopods CW = 108 (Haig 1974)

(del Solar 1972)

Previous generic names and

aldwyn 1985b

Macpherson 1988c, Sandberg and abdominal asymmetry McLaughlin 1998

Macpherson 1988c

Campodonico and Guzman 1972, Macpherson 1988c

Dawson and Female with reversed Y

map Distribution

aldwyn 1985b)

Drawing (Makarov 1962, Sandberg and McLaughlin 1998), drawings and photos (Macpherson 1988c)

Photo (Macpherson 1988a) Photos and drawings (Macpherson 1988c)

Drawings (Perez 1934, Macpherson 1988c) photos (Campodonico and Guzman 1972, Macpherson 1988b,c)

Photo and drawings (Sakai 1978), drawings (Dawson and Y

Photo (del Solar 1981)

or drawings

able 4. (Continued.)

Scientific and Locations of vernacular photographs names L. maja, northern stone crab, prickly crab

L. mamillifer L. manningi

L. murrayi

L. nintokuae

L. panamensis 782 Zaklan — Review of Lithodidae

L. longispina

Lithodes confundens (Macpherson 1988c) L. murrayi (Campodonico and Guzman 1972)

from Dawson 1989.

species

size (mm) related

Maximum Closely male/female

Male: CL = 102, et al. 1996) (Vinuesa CW = 102 (Macpherson 1990) (Macpherson 1990) (Boschi et al. 1984), female: CL = 142,

(CW = 65 Campodonico

of Male: CL = 198, CW = 250

of CW = 140 (Macpherson

comments

inogradov 1972

Only males found, L. antarcticus dedicated to Dr (Dawson 1989) Bertrand Richer of Jacquinot 1852 Pseudolithodes Forges of Orstom Named after Male: CL = 109, CW = 110, (Macpherson 1990) zenkevitschi Birstein and M. TurkayDr. 1988c), up to 8 kg female: CL = 66, V of Senckenberg Museum in and Guzman 1972) Frankfurt, Germany

(Macpherson 1988c)

Previous generic names and

Macpherson 1988c

map Distribution

Photo (Macpherson 1990) (Macpherson 1988c) Photo and drawings (Macpherson 1988c)

or drawings

centolla Photo and drawings

able 4. (Continued.) Photographs, distributions, maximum sizes, and general information concerning lithodids.

CL = carapace length (includes rostral length unless otherwise noted), CW = carapace width, R = rostrum length.

Macpherson 1988c maps are for Atlantic Ocean distributions only

For an exhaustive list see Dawson 1989. All uncited common Japanese names are obtained from Sakai 1976; other common names are

As a complete taxonomic account of each species is beyond the scope of this paper only selected junior synonyms are listed.

Scientific and Locations of vernacular photographs names

L. richeri

L. turkayi

L. santolla,

a b c d Crabs in Cold Water Regions: Biology, Management, and Economics 783

and

Closely

L. longispina (Wu et al. 1998) (Wu

L. murrayi (Macpherson

L. murrayi L. tropicalis

species

> 300

size (mm) related

Maximum male/female

CW = 155, female: CL = 133.5, CW = 132.1 (Wu et al. 1998) (Wu

Male: CL = 129, CW = 124, female: CL = 118, CW = 119 1988c) (Macpherson 1984)

Female: CL = 103.5, R = 16, CW = 97

CW = 128 (Haig 1974)

Male: CL = 185, CW = 165, female: CL = 175, (Jensen 1995) CW = 145 (Hart 1982)

Male: CL = 200, CW = 270 (Hart 1982), CW

, fide Male: CL = 101, R = 9,

Echinocerus

comments

Paralomis

Females only and found on trawler “Wiracocha” (creator“Wiracocha” (Haig 1974) (Haig 1974) God in Inca myth, Haig 1974)

In Macpherson 1988c

Includes White (Dawson 1989)

Previous generic names and

Sakai 1976 Male: CL = 149.0,

Macpherson 1988c

map Distribution

et al. 1998), drawings

Wu (Sakai 1976)

Photos (Macpherson 1984), drawings (Macpherson 1988c)

Drawings (Haig 1974)

Photo (del Solar 1981)

Drawings (Hart 1982, 1921, Jensen 1995) (Schmitt 1921, Jensen 1995)

or drawings

Ibaragani Photo (Sakai 1971, 1976,

, box crab, Drawing (Makarov 1962,

able 4. (Continued.)

Scientific and Locations of vernacular photographs names

L. turritus,

L. unicornis

L. wiracocha

Lopholithodes diomedeae

L. foraminatus, box crab Macpherson 1988c), photo

L. mandtii noduled crab Hart 1982), photo (Schmitt 784 Zaklan — Review of Lithodidae

Closely

N. agassizii (Macpherson 1988c)

from Dawson 1989.

species

size (mm) related

Maximum male/female

(Sakai 1980)

Male: CL = 167, CW = 162,

CW = 143 (Macpherson 1988c)

Male: CL = 195, CW = 189, female: CL = 180, CW = 156

(Macpherson 1988c)

. T. Male: CL = 41, CW = 47

Barnard

aff.

L. agassizii

Paralomis

comments

Named after Dr Odawara, director of the Odawara Carcinological Museum, Tokyo (Sakai Museum, Tokyo 1980) in fide Macpherson 1988c

Lithodes agassizii (Smith 1882)“Investigator” authors female: CL = 154, and (?) asperrimus of Macpherson 1988 Includes (Dawson 1989)

Previous generic names and

Macpherson 1988c

map Distribution

Drawings (Smith 1882, Macpherson 1988c)

Photo (Macpherson 1983, Macpherson 1988a), drawings and photos (Macpherson 1988c)

or drawings

able 4. (Continued.) Photographs, distributions, maximum sizes, and general information concerning lithodids.

CL = carapace length (includes rostral length unless otherwise noted), CW = carapace width, R = rostrum length.

Macpherson 1988c maps are for Atlantic Ocean distributions only

For an exhaustive list see Dawson 1989. All uncited common Japanese names are obtained from Sakai 1976; other common names are

As a complete taxonomic account of each species is beyond the scope of this paper only selected junior synonyms are listed.

Scientific and Locations of vernacular photographs names L. odawarai

Neolithodes

Neolithodes agassizii

N. alcocki N. asperrimus

a b c d Crabs in Cold Water Regions: Biology, Management, and Economics 785

and

Closely

aldwyn 1970)

N. agassizii N. asperrimus

N. vinogradovi N. grimaldii (Macpherson 1988c)

N. agassizii, N. capensis N. asperrimus

(Sakai 1976)

species

size (mm) related

Maximum male/female

CL = 130, R = 18, CW = 104 (Dawson and Yaldwyn 1970)and Yaldwyn (Dawson and

Male: CL = 131, CW = 130, female: CL = 86, CW = 86 (Macpherson 1988c)

Male: CL = 197,

CL = 41.5, CW = 35.0 (Baez et al. 1986)

Male: CL = 152,

CL = 153, CW = 138

Male: CL = 168, CW = 142 (Sakai 1976)

comments

inogradov 1972

Named after J.W Brodie, Director of the New Zealand Oceanographic Institute (Dawson and Yaldwyn 1970) and Yaldwyn

Neolithodes martii Birstein and CW = 166, female: V (Macpherson 1988c)

Lithodes goodei

Lithodes agassizii Smith 1882 (Macpherson 1988c) (Dawson 1989)

Previous generic names and

Macpherson 1988c

Macpherson 1988c, Sandberg and Benedict 1895 CW = 145, female: McLaughlin 1998

map Distribution

Photo and drawings (Macpherson 1988c)

Drawings (Baez et al. 1986, Macpherson 1988c)

Photo and drawings

and McLaughlin 1998)

Drawings (Sakai 1971, 1976), photo (Miyake 1982)

or drawings

able 4. (Continued.)

Scientific and Locations of vernacular photographs names N. brodiei

N. capensis

N. diomedeae, centolla patache (Baez et al. 1986)

N. grimaldii, Atlantic porcupine stone crab (Macpherson 1988c), (Pohle 1992) drawing (Sandberg

N. nipponensis, nihon-ibaragani 786 Zaklan — Review of Lithodidae

platypus

N. grimaldii (Macpherson 1988c)

Can hybridize with P. (Nizyayev 1991)

from Dawson 1989.

species

size (mm) related

Maximum Closely male/female

female: CL = 91, CW = 105 (Macpherson 1990)

Male: CL = 95, CW = 102 92

CL = 118 (Sato and Abe 1941)

Male: CL = 227, CW = 283, female: CL = 195,CW = 213 (Powell and Nickerson

1965b)

rostro-

L. spino-

of MacKay

of Brandt 1848

comments

Up to 11 kg body (Hart 1982), up to 21 years (Matsuura and Takeshita 985), and Takeshita

and Munk 1991)

falcatus 1932 and sissimus (Dawson 1989), gynandromorphistic individual (Stevens

Previous generic names and

. L.G. Male: CL = 109, CW = 113

Named after Dr

(Macpherson 1988c)

Sakai 1971, 1976; Abe 1992 (Sakai 1976), female:

Sakai 1976, Otto et al. 1980, Abe 1992, Sandberg and McLaughlin 1998 1 includes

map Distribution

Photos (Macpherson 1988c,

1990) Vinogradov (Macpherson 1988c),

Sakai 1976

Drawing (Makarov 1962,

Sakai 1971, 1976),

Photo (Schmitt 1921)

Drawing (Makarov 1962,

Sandberg and McLaughlin 1998), photo (Miyake 1982,

Munk 1991, Jensen 1995), drawing (Macpherson 1988c)

or drawings

able 4. (Continued.) Photographs, distributions, maximum sizes, and general information concerning lithodids.

brevipes,

californiensis

camtschaticus,

arabagani, Sakai 1971, 1976, Hart 1982,

CL = carapace length (includes rostral length unless otherwise noted), CW = carapace width, R = rostrum length.

Macpherson 1988c maps are for Atlantic Ocean distributions only

For an exhaustive list see Dawson 1989. All uncited common Japanese names are obtained from Sakai 1976; other common names are

As a complete taxonomic account of each species is beyond the scope of this paper only selected junior synonyms are listed.

Scientific and Locations of vernacular photographs names N. vinogradovi

Paralithodes

P. hanasaki crab photo (Miyake 1982) Hanasaki-gani, (Abe 1992)

P. T Alaskan, Russian, Japanese or red king crab Bliss 1983, Stevens and

a b c d Crabs in Cold Water Regions: Biology, Management, and Economics 787

Closely

camtschaticus

indica

cristulata, P.

africana

Can hybridize with P.

(Nizyayev 1991)

P. (Macpherson 1982), P. anamerae (Macpherson 1988c)

P. (Macpherson

species

size (mm) related

Maximum male/female

CL = 159, CW = 170 (Sakai 1976)

Male: CL = 79, female: CL = 68 (Macpherson 1988c)

Male: CL = 97, CW = 102, female: CL = 68, CW = 68 1988c) (Macpherson 1988c)

and

in Benedict

comments

rawlers Owners

Up to 17 years (Jensen and Armstrong 1989)

Leptolithodes Pristopus

Acantholithus Stimpson 1859 (Dawson 1989)

Named after the Asociacion Nacional de Armadores de Buques Congeladores de Pesca de Merluza (ANAMER, Hake Macpherson 1988c) Fishery Freezer T National Association;

Previous generic names and

akeda 1974, Synonymous with

Sakai 1976, Otto et al. 1980, Abe 1992

T Sakai 1971, Macpherson 1988c 1895 and (?)

Macpherson 1988c

map Distribution

Drawing (Makarov 1962,

photo (Miyake 1982)

Photo (Schmitt 1921)

Photo and drawings (Macpherson 1982), drawings (Macpherson 1983, Macpherson 1988c)

Photos and drawings (Macpherson 1988c)

or drawings

able 4. (Continued.)

platypus,

rathbuni

africana

anamerae

Scientific and Locations of vernacular photographs names P. Aburagani, Sakai 1971, 1976, Sandberg blue king crab and McLaughlin 1998),

Paralomis

P. 788 Zaklan — Review of Lithodidae

Closely

spectabilis

investigatoris

P. (Macpherson

P. (Macpherson 1989)

Glyptolithodes

1979)

from Dawson 1989.

species

size (mm) related

Maximum male/female

CL = 75, CW = 75 (Hart 1974)

CW = 53 1988b) (Macpherson 1988b)

Male: CL = 78, CW = 74, female: CL = 55,

Male: CL = 23, CW = 21,

CW = 34 (Hansen 1908)

Male: CL = 52, CW = 57 (Macpherson 1989)

Male: CL = 89, R = 11, CW = 96 (Takeda andCW = 96 (Takeda and Ohta (Takeda Ohta 1979), CL = 108

(Sakai 1987)

. Y.

spectabilis

comments

inogradov 1967,

not of Hansen 1908 (Dawson 1989); named after Dr of Birstein and Birstein from State V of Moscow Univ. (Macpherson 1988b)

Previous generic names and

inogradov 1967

Birstein and Includes

Macpherson 1988c, Sandberg and female: CL = 34.8, McLaughlin 1998

map Distribution

Photo (del Solar 1981)

Photo and drawings (Birstein

and Vinogradov 1967, and Vinogradov Macpherson 1988b)

Drawings (Hansen 1908, Macpherson 1988c, Sandberg and McLaughlin 1998), photos (Pohle 1992)

or drawings

, Hiraashi and Ohta 1979) Photos (Takeda

ceres

able 4. (Continued.) Photographs, distributions, maximum sizes, and general information concerning lithodids.

aspera

birsteini

bouvieri

cristata

CL = carapace length (includes rostral length unless otherwise noted), CW = carapace width, R = rostrum length.

Macpherson 1988c maps are for Atlantic Ocean distributions only

For an exhaustive list see Dawson 1989. All uncited common Japanese names are obtained from Sakai 1976; other common names are

As a complete taxonomic account of each species is beyond the scope of this paper only selected junior synonyms are listed.

Scientific and Locations of vernacular photographs names P.

P. -ezoibaragani (Takeda and (Takeda

Ohta 1979) a b c d Crabs in Cold Water Regions: Biology, Management, and Economics 789

(Andrade

Closely

aspera

investigatoris

zealandica

P. 1980)

P. (Chace 1939)

P. (Feldmann 1998)

species

size (mm) related

Maximum male/female

Male: CL = 47.2, CW = 47.6, female: CL = 55.4, CW = 56.4 (Andrade 1980)

(Macpherson 1988c)

Female: CL = 61.2, CW = 53.0 (Chace 1939) (Macpherson 1988c) CL = 94, CW = 99 (Sakai 1976)

” small Female: CL = 55, CW = 57

” urchin Female: CL = 78, CW = 76

cristula

ericius

comments

Latin “ crest, referring to crests on walking legs and lateral carapace edges (Macpherson 1988c)

or hedgehog, due to the spiny carapace, only female specimens found (Macpherson Latin “

1988c)

Previous generic names and

Macpherson 1988c

Anne Debode Macpherson 1988c (Feldmann 1998)

Fossil only, namedFossil only, Sakai 1971, 1976 CL = 63.6, CW = 53.2 after John and (Feldmann 1998) Macpherson 1988c

map Distribution

Drawings (Andrade 1980)

Drawings (Macpherson 1988c)

Photos and drawings (Macpherson 1988c) (Macpherson 1988c)

Photo and drawing Drawing (Balss 1911), photo (Feldmann 1998) (Sakai 1971, 1976) Photo and drawings

or drawings

able 4. (Continued.)

chilensis

cristulata

cubensis

debodeorum dofleini,

erinacea

subu-ezoibaragani

Scientific and Locations of vernacular photographs names P.

P. P. T

P. 790 Zaklan — Review of Lithodidae

and

aspera,

and

(Yumao et (Yumao

(Sakai

(Eldredge

Closely

inca

spectabilis

dofleini

longidactyla papillata

dofleini,

hystrix papillata, hystrix

P. (Macpherson 1988c)

P. P. (Macpherson 1988c)

1976)

P. P. al. 1984) P. P. 1980)

from Dawson 1989.

species

size (mm) related

Maximum

male/female

Male: CL = 95, CW = 96, female: CL = 89, CW = 87 (Macpherson 1988c)

CW = 95 (Takeda and CW = 95 (Takeda Hatanaka 1984), CL = 1201.5 kg max (Vinuesa et al. 1996) (Vinuesa

Female: CL = 94, CW = 106

Male: CL = 95, CW = 95,

(Eldredge 1976)

CL = 50, CW = 55 female: CL = 97, et al. 1984) (Yumao CW = 93.5 Male: CL = 81, CW = 78 (Sakai 1980), CL = 70-95

(Sakai 1987)

comments

Only species of this genus that inhabits shallow coastal waters (Macpherson 1988c)

Only females known, named after G.D. Grossman (Macpherson 1988c) (Macpherson 1988c)

Previous generic names and

Macpherson 1988c

Distribution map

Photo (Macpherson 1988c)

Photo (Campodonico 1978, Ingle and Garrod 1987),

(Macpherson 1988c)

Photo (Eldredge 1976,

Macpherson 1990) Photo (Sakai 1987)

or drawings

able 4. (Continued.) Photographs, distributions, maximum sizes, and general information concerning lithodids.

formosa

granulosa,

grossmani

haigae

heterotuberculata

hystrixoides

CL = carapace length (includes rostral length unless otherwise noted), CW = carapace width, R = rostrum length.

Macpherson 1988c maps are for Atlantic Ocean distributions only

For an exhaustive list see Dawson 1989. All uncited common Japanese names are obtained from Sakai 1976; other common names are

As a complete taxonomic account of each species is beyond the scope of this paper only selected junior synonyms are listed.

Scientific and Locations of vernacular photographs

names P.

P. false king crab, centollon photos and drawings

a b c d Crabs in Cold Water Regions: Biology, Management, and Economics 791

(Alcock

(Takeda

(Alcock and

Closely

multispina

verrucosa

aspera

africana

P. (Sakai 1976)

P. and Anderson

P. Anderson 1899)

P. and Hashimoto 1990)

species

size (mm) related

Maximum male/female

CW = 114 (Sakai 1976)

Male: CL = 80, R = 10.5, CW = 93, female: CL = 108, R = 13, CW = 123 (Haig 1974)

CL = 39.5, CW = 37 (Alcock and Anderson 1899) 1899)

CL = 33, CW = 29.5 (Alcock and Anderson 1899)

CL = 39, CW = 36 (Sakai 1971)

Male: CL = 64.7, CW = 67.8, female: CL = 70.5, CW = 68.8

1990)

CL = 66.2, CW = 61.9

comments

echnology Center

Marine Science T (JAMSTEC; Takeda and Hashimoto (Takeda and Hashimoto 1990)

Kyushu-Palau 1985) (Takeda submarine ridge

(Takeda 1985) (Takeda

Previous generic names and

Sakai 1976 CL = 106, R = 20,

Sakai 1971, 1976

map Distribution

Photo (Sakai 1971, 1976,

Photos (Haig 1974, del Solar 1981)

Drawings (Balss 1911, Sakai 1971, 1976), photo (Miyake 1982)

Photos (Takeda andPhotos (Takeda Named after Japan Hashimoto 1990)

Photo (Takeda 1985)Photo (Takeda Named after the

or drawings

able 4. (Continued.)

hystrix,

inca

indica

investigatoris

japonica,

jamsteci,

kyushupalauensis

Scientific and Locations of vernacular photographs names P. Igagurigani Miyake 1982)

P. Kofuki-ezoibaragani

P. Ensei-ezo-ibaragani (Takeda and (Takeda Hashimoto 1990)

P. 792 Zaklan — Review of Lithodidae

(Williams

Closely

grossmani

pentinata serrata cristulata africana

P. (Macpherson 1988c)

P. P. P. P. et al. 2000)

from Dawson 1989.

species

size (mm) related

Maximum male/female

Male: CL = 99, CW = 105 (Macpherson 1988c)

CL = 106, R = 14, CW = 117 (Haig 1974)

Male: CL = 146, CW = 133 (Williams

(Takeda 1974) (Takeda

Female: CL = 15, CW = 15 (Macpherson 1988c)

CL = 80, CW = 78 (Benedict 1895), male: CL = 76, CW = 82 (Sakai 1971)

comments

Only one male specimen (Birstein and Vinogradov 1972) and Vinogradov

Manning (Williams et al. 2000) et al. 2000)

Only one specimen female (Macpherson 1988c)

Previous generic names and

Macpherson 1988c

Sakai 1971, 1976

map Distribution

inogradov 1972), photo

Drawings (Birstein and V and drawings (Macpherson 1988c)

Drawings (Haig 1974), photo (del Solar 1981)

Drawings (Williams Only males known, et al. 2000) named after R.B.

Photos (Takeda 1974)Photos (Takeda CL = 70, CW = 58

Dorsal and drawings (Macpherson 1988c)

Drawing (Makarov 1962, Hart 1982), photo (Schmitt 1921, Sakai 1971, 1976)

or drawings

able 4. (Continued.) Photographs, distributions, maximum sizes, and general information concerning lithodids.

longidactyla

longipes

manningi

medipacifica

microps

multispina,

CL = carapace length (includes rostral length unless otherwise noted), CW = carapace width, R = rostrum length.

Macpherson 1988c maps are for Atlantic Ocean distributions only

For an exhaustive list see Dawson 1989. All uncited common Japanese names are obtained from Sakai 1976; other common names are

As a complete taxonomic account of each species is beyond the scope of this paper only selected junior synonyms are listed.

Scientific and Locations of vernacular photographs names P.

P. Ezo-ibaragani

a b c d Crabs in Cold Water Regions: Biology, Management, and Economics 793

(Sakai

seagranti

Closely

zealandica

serrata

spinossima

investigatoris

P. 1978)

P. (Macpherson

P. (Macpherson 1992)

P. and

species

size (mm) related

Maximum male/female

Male: CL = 72 , CW = 78

CW = 83.5, female: CL = 98.3, CW = 96.0 (Wilson 1990) (Wilson

Male: CL = 74, R = 8,

Male: CL = 118, R = 10, CW = 130 (Haig 1974)

Female: CL = 50, CW = 45 (Kensley 1981)

” comb, Female: CL = 96, CW = 96

pectinata

comments

From Greek “ochthos” hilly elevation (Macpherson 1988b) (Macpherson 1988b)

Latin “ referring to (Macpherson 1988c) pereiopod 1988c) spinulation; only one female (Macpherson 1988c)

Greek “phrixos” bristled (Macpherson 1992)

Named after Martina Roeleveld (Kensley

1981) (Kensley 1981)

Previous generic names and

Only one male known (Sakai 1978) CW = 77 (Sakai 1978)

Macpherson 1988c

map Distribution

Dorsal and photos (Macpherson 1988b)

Drawings (Wilson 1990)Drawings (Wilson Male: CL = 84.6,

Photo and drawings

Drawings (Haig 1974), photo (del Solar 1981)

Photos (Macpherson 1988c)

Photos (Macpherson 1992)

Photo and drawings (Kensley 1981)

or drawings

able 4. (Continued.)

ochthodes

otsuae

pacifica

papillata

pectinata

phrixa

roeleveldae

Scientific and Locations of vernacular photographs names P.

P. 794 Zaklan — Review of Lithodidae

hystrix-

and

inca

, and

erinacea

aspera, P. verrilli, aspera, P. longipes, P. inves- longipes, P.

pectinata verrilli

hystrix

spinosissimus

granulosa

P. P. tigatoris, P. medi tigatoris, P. pacifica juvenile (Eldredge 1976

P. P. (Macpherson 1988c)

P. oides P. bouvieri oides P. P. Hatanaka 1984), and (Macpherson 1988c)

P. (Macpherson 1990)

from Dawson 1989.

species

size (mm) related

Maximum Closely male/female

Male: CL = 80.5, CW = 81,

CW = 57 (Eldredge 1976)

(Macpherson 1988c)

Female: CL = 75, CW = 78 (Takeda and Hatanaka (Takeda 1984)

Female: CL = 71, CW = 72 (Macpherson 1990)

” saw, due” saw, Male: CL = 106, CW = 112

serra

comments

ebber and Dawson at

Named after the office of Sea Grant female: CL = 62, Programs (Eldredge 1976)

Latin “ to pereiopod spines; only one male (Macpherson 1988c)

Presently studied by W the National Museum 1990) of New Zealand; no males (Macpherson

Previous generic names and

Macpherson 1988c

map Distribution

Photos (Eldredge 1976)

Photos and drawings (Macpherson 1988c)

Photo and drawings (Takeda and Hatanaka 1984, (Takeda Macpherson 1988c)

Photo and drawings (Macpherson 1990)

or drawings

sp.

able 4. (Continued.) Photographs, distributions, maximum sizes, and general information concerning lithodids.

seagranti

serrata

shinkaimaruae

CL = carapace length (includes rostral length unless otherwise noted), CW = carapace width, R = rostrum length.

Macpherson 1988c maps are for Atlantic Ocean distributions only

For an exhaustive list see Dawson 1989. All uncited common Japanese names are obtained from Sakai 1976; other common names are

As a complete taxonomic account of each species is beyond the scope of this paper only selected junior synonyms are listed.

Scientific and Locations of vernacular photographs names P.

(Takeda and (Takeda

a b c d Crabs in Cold Water Regions: Biology, Management, and Economics 795

Closely

birsteini shinkaimaruae formosa

indica erinacea,

granulosa

P. P. P. (Macpherson (Macpherson 1988c) 1988c) P. P. (Macpherson 1988a)

P. (Macpherson 1988b)

species

inogradov

size (mm) related

Maximum male/female

Male: CL = 90, CW = 78 MacIntosh 1996), female: CL = 100, CW = 103 1967), female: CL = 45.5, (Macpherson 1988c) CW = 46 (Hansen 1908) Male: CL = 71, CW = 72, Male: CL = 125 (Otto and female: CL = 49, CW = 50 (Macpherson 1988a)

Male: CL = 45, CW = 40, female: CL = 45, CW = 39 (Takeda and Miyake 1980) (Takeda

CL = 41, CW = 47

CL = 102, R = 10, CW = 113 (Sakai 1971), male: CL = 112,

CW = 102 (Hart 1982)

comments

Latin “tuber” tubercle, and “pes” foot (Macpherson 1988b) (Macpherson1988b)

Previous generic names and

Macpherson 1988c, Sandberg and (Birstein and V McLaughlin 1998

Macpherson 1988c

Sakai 1971, 1976

map Distribution

inogradov 1972), photo and

Drawings (Hansen 1908, Sandberg and McLaughlin drawings (Macpherson 1988c) 1998), photos and drawings (Macpherson 1988c) Photo and drawings Drawings (Birstein and (Macpherson 1988a) V

Photo (Takeda and Miyake Photo (Takeda 1980)

Photo and drawings (Macpherson 1988b)

1921, Sakai 1971, 1976,

1987, Miyake 1982)

or drawings

, Gokaku- Drawing (Makarov 1962,

rrilli

able 4. (Continued.)

spectabilis

stella spinosissima

truncatispinosa,

tuberipes

Scientific and Locations of vernacular photographs names P.

P. P.

P. Ibo-ezoibaragani (Takeda and (Takeda Miyake 1980)

P. ezoibaragani Hart 1982), photo (Schmitt 796 Zaklan — Review of Lithodidae

Closely

from Dawson 1989.

species

aldwyn

size (mm) related

Maximum male/female

Male: CL = 111, CW = 101 (Dawson and Y

1971)

Male: CL = 90, CW = 90, female: CL = 50, CW = 60 (Hart 1982)

Male: CL = 59, CW = 64, female: CL = 50, CW = 57 (Hart 1982)

Male: CL = 38, CW = 36, female: CL = 32, CW = 31 (Makarov 1934)

(Hart 1982)

Male: CL = 62, CW = 64, female: CL = 49, CW = 50

comments

and Yaldwyn 1971) and Yaldwyn

Previous generic names and

map Distribution

aldwyn Named after New

Photos (Dawson and Y 1971) Zealand (Dawson

Drawing (Makarov 1962, Hart 1982, Macpherson 1988c), photos (Schmitt 1921, Jensen 1995)

Photo (Makarov 1934), drawing (Makarov 1962, Sakai 1971, 1976, Macpherson 1988c)

Drawing (Hart 1982), photo (Schmitt 1921, Jensen 1995)

or drawings

able 4. (Continued.) Photographs, distributions, maximum sizes, and general information concerning lithodids.

zealandica

CL = carapace length (includes rostral length unless otherwise noted), CW = carapace width, R = rostrum length.

Macpherson 1988c maps are for Atlantic Ocean distributions only

For an exhaustive list see Dawson 1989. All uncited common Japanese names are obtained from Sakai 1976; other common names are

As a complete taxonomic account of each species is beyond the scope of this paper only selected junior synonyms are listed.

Scientific and Locations of vernacular photographs names P.

Phyllolithodes papillosus

Rhinolithodes wosnessenskii, rhinoceros crab

Sculptolithodes derjugini, Eri-tarabagani

Subfamily Hapalogastrinae Acantholithodes a b hispidus c d Crabs in Cold Water Regions: Biology, Management, and Economics 797

species

size (mm) related

Maximum Closely male/female

CL = 23.0, CW = 23.0 (Makarov 1962), male: CL = 25.0, CW = 21.0, female: CL = 11.20, CW = 9.78 (Zaklan, pers. obs.)

CL = 20 (Jensen 1995), female: CL = 12.62, CW = 14.71 (Zaklan, pers. obs.)

Male: CL = 21.5, female: CL = 15.6 (Goshima et al. 1995)

CL = 20.5, CW = 22 (Miyake 1982), male: CL = 23, CW = 24 (Hart 1982)

Male: CL = 25, CW = 25, female: CL = 22, CW = 24 (Hart 1982), CL = 35 (Jensen 1995)

Male: CL = 25, CW = 30, female: CL = 22,

of CW = 20 (Hart 1982)

of Benedict

comments

Hapalogaster inermis of Stimpson 1860, H. brandtii Schalfeew 1892, O. gilli

1895 (Dawson 1989)

Previous generic names and

map Distribution

Drawing (Schalfeew 1892,

(Miyake 1982, Jensen 1995)

Photo (Schmitt 1921, Jensen 1995)

Drawing (Makarov 1962), photo (Miyake 1982)

Drawing (Schalfeew 1892,

photo (Schmitt 1921, Miyake 1982)

Drawing (Schalfeew 1892, Hart 1982), photo (Jensen 1995)

Drawing (Schalfeew 1892,

photo (Schmitt 1921, Miyake 1982, Jensen 1995)

or drawings

, Ibo-gani, Makarov 1962, Hart 1982),

able 4. (Continued.)

Scientific and Locations of vernacular photographs names Dermaturus mandtii, hairy crab Makarov 1962), photo

Hapalogaster cavicauda, furry crab

Hapalogaster dentata

H. grebnitzkii, Syojo-gani Makarov 1962, Hart 1982),

H. mertensii

Oedignathus inermis 798 Zaklan — Review of Lithodidae

from Dawson 1989.

species

size (mm) related

Maximum Closely

male/female

Male: CL = 61.5,

1892), female: CL = 50, CW = 53 (Hart 1982)

Lepeopus

wosnessenskii

comments

Benedict 1895 CW = 73.3 (Schalfeew (Dawson 1989)

Includes Probably synonymous with

(Dawson 1989)

Previous generic names and

Distribution map

Makarov 1962, Hart 1982),

Jensen 1995)

Drawing (Schalfeew 1892,

or drawings

able 4. (Continued.) Photographs, distributions, maximum sizes, and general information concerning lithodids.

CL = carapace length (includes rostral length unless otherwise noted), CW = carapace width, R = rostrum length.

Macpherson 1988c maps are for Atlantic Ocean distributions only

For an exhaustive list see Dawson 1989. All uncited common Japanese names are obtained from Sakai 1976; other common names are

As a complete taxonomic account of each species is beyond the scope of this paper only selected junior synonyms are listed.

Scientific and Locations of vernacular photographs names wosnessenskii, Urokogani photo (Sakai 1971, 1976,

Placetron Placetron forcipatus

a b c d

Crabs in Cold Water Regions: Biology, Management, and Economics 799

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile, Size of first of Size

2.0 CW

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

(Z) glaucothoe (Z) Size of of zoea of of Size

G = 2.5 G = 10.5 (Kim (Kim and and Hong Hong 2000) 2000)

(Hart 1965)

stages

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal Number of Number

(Kim and Z2 = 2.2 Z2 = 8.9 (Kim and

2000) Z4 = 3.2 Z4 = 14.8

Eclosion date Eclosion

(mm)

Egg diameter Egg

(eggs/clutch)

Brood size Brood

cycle Reproductive

pers. obs.) obs.)

(Hart 1965) 1965) G = 1.6 CW

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 25.84, 4Z, 1G Z1 = 1.6 Z1 = 4.8 Z1-4 and G R = 5.88, CW = 35.82 Hong Z3 = 2.6 Z3 = 8.2 Hong 2000) (Zaklan, pers. obs.)

CL = 35.97, Uneyed 471 1.01- CW = 49.84 embryos in (Zaklan, 1.06 (Zaklan, August pers. obs.) (Zaklan, pers. obs.) (Zaklan, pers.

CL = 24.5, Uneyed eggs 0.75-0.92 March- PZ, 4 Z, Z1 = 3 TL Z1-Z4 = C1 = Z1-4 and G CW = 36; in August and just April 1 G Z2 = 3.25 TL 14-16 2.25 CL and C1 (Hart possibly with de- before (Hart (Hart Z3 = 3.4 TL (Hart= C1 1965) CL = 18.5, velopmental hatching 1965) 1965) Z4 = 3.6 TL 1965) CW = 26.5 signs in 0.8-1.1 = 2.8 TL G (Hart 1965) (Hart 1965) December (Hart G = 1.9

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size

Age (years) and (years) Age able 5. Life history traits of the family Lithodidae. Species

Subfamily Lithodinae Cryptolithodes expansus

C. sitchensis

C. typicus 800 Zaklan — Review of Lithodidae

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile, Size of first of Size

Acantholithodes hispidus

(days)

cothoe stages cothoe

zoeal and glau- and zoeal Duration of Duration

dent, Paul and Paul 1999)

(G) (mm) (G)

a (Z) glaucothoe (Z)

juvenile instar ( = crab one); C2 = second juvenile Size of of zoea of of Size

Cryptolithodes sitchensis

. stages

glaucothoe (G) glaucothoe zoeal (Z) and (Z) zoeal

and G Z1 = 7.3 TL Z1 = 6.6 Z 1-4 and G

Number of Number Eclosion date Eclosion

Phyllolithodes papillosus

Oedignathus inermis

February- 4 Z

1997) depen-

(mm)

Egg diameter Egg

(eggs/clutch) Brood size Brood

2,600-5,500 2.3 (Somer-

cycle Reproductive

1981) (Somerton

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size

Age (years) and (years) Age

turity of males of turity productive ma- productive

(courtesy of Greig Jensen, C. Nyblade, and Hokkaido University), and size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 114 CL = 105 Spawn July- 9,500- 2.1 (Hira- based on (Japan, Hira- October 30,100 moto and July Haynes Z2 = 7.5 TL Z2 = 7-8.8 (Haynes chelae moto and (Hiramoto (Hiramoto Sato (Hira- 1982), 3 Z Z3 = 7.6 TL Z3 = 12 1982) allometry Sato 1970, and Sato and Sato 1970), 2.4 moto skips Z3 or Z4 = 6.8 TL Z4 = 9.3 (northern B.C., Canada, 1970), 1970) (Jewett and Sato Z4 (Shirley G = 5.9 TL G = 41.3 B.C., Jewett Jewett et al. aseasonal et al. 1970), and Zhou (Haynes Z1-4 and et al. 1985), 1985), reproduction 1985), March 1997), 3 Z 1982 G = 67 CL = 92-130 CL = 98-111 (Sloan 1985, 2.07-2.52 (Haynes (Paul and (Shirley (latitude (latitude de- Somerton (Zaklan, 1982), Paul 1999, and Zhou dependent; pendent; and Otto pers. obs.) April- Adams and 1997), Somerton Somerton 1986) August Paul 1999) Z1-4 and and Otto and Otto (Shirley G = 75- 1986), 1986) and Zhou 148 (Tº functional CL = 107 (Paul and Paul 2001)

CL = 91.4 CL = 80.2 Asynchronous (Somerton (Somerton or protracted (Somerton ton 1981) 1981) 1981) spawning 1981)

able 5. (Continued.) Life history traits of the family Lithodidae. Species Measurement of carapace length (CL) unless otherwise noted.

T Lithodes aequispinus

L. couesi

R = rostrum; CL = carapace length; CW = carapace width; TL = total length; G = glaucothoe; Z = zoea; PZ = pre-zoea; C1 = first instar ( = crab two). McLaughlin and Lemaitre (2000) report the possession of glaucothoeal and first crab stages of Hapalogaster dentata a

Crabs in Cold Water Regions: Biology, Management, and Economics 801

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile,

Size of first of Size

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

(Z) glaucothoe (Z) Size of of zoea of of Size

4.3 TL

stages

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal

Number of Number

Eclosion date Eclosion

(mm)

Egg diameter Egg 1.8 2 Z, 1 G Z1 = Z1 = 4 C1 = Z1-2 and G

(eggs/clutch)

Brood size Brood

cycle Reproductive

(Hiramoto 1974) 1974)

aseasonal (CL = 2.7-2.4) hatching of (Macdonald zoea (Anger et al. 1957) 1996)

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 98 Spawning 3,900- (Hiramoto August- 11,200 1974) October (Hiramoto

CL = 59, Spawning 2.0 CW = 57 September- (Macdonald (Macdonald 6.7-8.5 TL Z2 = 5 2,759 mg (Macdonald et (Macpherson November, et al. 1957) et al. 1957) (CL = 4.2-5.1) Z3 = 8 (Anger al. 1957) 1988c), zoeal 3 Z, 1 G Z2 = 6.7- G = 29 1996) CL = 61.16, eclosion in (Anger 9.0 TL C1 = 31 R = 21.53, April/May 1996) (CL = 4.2- (Anger CW = 56.92 (Pike and 5.4), 1996) (Zaklan, Williamson G = 4.6- pers. obs.) 1959)

CL = 123, 1988a) CW = 118 (Macpherson

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 108 CL = 69 Aseasonal 8,000 1.97 (Abello and (Macpherson (Abello and (Abello and (Abello and Macpherson 1988c) Macpherson Macpherson Macpher- 1992) 1992) 1992) son 1992) able 5. (Continued.) Species

L. ferox

L. longispina

L. maja

L. mamillifer 802 Zaklan — Review of Lithodidae

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile, Size of first of Size

Acantholithodes hispidus

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

(Z) glaucothoe (Z) Size of of zoea of of Size juvenile instar ( = crab one); C2 = second juvenile

Cryptolithodes sitchensis

stages .

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal

Number of Number Eclosion date Eclosion

Phyllolithodes papillosus

Oedignathus inermis

(mm)

Egg diameter Egg

(eggs/clutch) Brood size Brood

and Do-Chi Do-Chi 1977)

cycle

Reproductive

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

aldwyn 1985b)

CW = 120 CL = 106 (Macpherson 1988c)

(Miquel et al. 1985)

CL = 62 (Dawson and Y

CL-R = 100 (Haig 1974) CW = 108

turity of males of turity productive ma- productive

(courtesy of Greig Jensen, C. Nyblade, and Hokkaido University), and size (mm) at re- at (mm) size Age (years) and (years) Age

Chela CL = 60-66 380-3,582 1.92-2.88 allometry (Arnaud and (Arnaud (Arnaud and CL = 69.5- Do-Chi 1977), 71.0 chela 1977) (Miquel et allometry al. 1985) CL = 64.5-65.5

able 5. (Continued.) Life history traits of the family Lithodidae. Species Measurement of carapace length (CL) unless otherwise noted.

L. manningi

L. murrayi

L. nintokuae

L. panamensis

Crabs in Cold Water Regions: Biology, Management, and Economics 803

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile,

Size of first of Size

(days)

cothoe stages cothoe

zoeal and glau- and zoeal Duration of Duration

inuesa

dent Campo- donico 1971, V et al. 1985)

(G) (mm) (G)

(Z) glaucothoe (Z)

Size of of zoea of of Size

stages

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal

Number of Number Eclosion date Eclosion

October donico Z2 = 2.06- Z3 = 14- 1992) (Campodoni-

(mm) Egg diameter Egg

and Campo-

1999)

(eggs/clutch) Brood size Brood

60,000 (Vinuesa 1999) 3.83 G = 33-

cycle Reproductive

inuesa 1987)

egg release donico donico (Lovrich 1971) 3.75 26 co 1971)

(Hernandez 1985, V

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 77.5 December to 4111 1.40-2.47 Mid-Sept- 3 Z, 1G Z1 = 1.88- Z1 = 4-12 C1 = 1.5 Z1-3 and G

1972), gonadal

CL = 58, 1.7 CW = 55 (Lovrich (Macpherson and 1988c) Vinuesa

CL = 123.7, CW = 127.7 (Wu et al. 1998) (Wu 1988c) CL = 118, 1.5 CW = 115 (Macpherson (Macpherson 1984)

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

Morphometric maturity (Guzman and mid-January (Guzman (Guzman ember- (Campo- 3.75 Z2 = 4-16 (Oyarzun drawings CL = 90-99 Campodonico molt/mate/ and Campo- (Boschi et al. 1984), maturity (Hernandez 1972), to 1972), 2.1 and Vinuesa Z3 = 2.06- Z and gonadal CL = 66-87 1985), annual maturity (Vinuesa 1991, (Vinuesa (Vinuesa 1987) G = 1.8- 55 (Tº CL = 60-75 1984), age 4, Lovrich and 1982) 2.16 and (Vinuesa oogenesis 1999), Vinuesa (Campo- salinity 1984), age 5 24 months embryo- donico 1971) depen- (Vinuesa et (Vinuesa genesis 1991) al. 1991) 9-11 months able 5. (Continued.) Species

L. santolla

L. turkayi

L. turritus

L. unicornis 804 Zaklan — Review of Lithodidae

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile, Size of first of Size

Acantholithodes hispidus

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

a (Z) glaucothoe (Z)

juvenile instar ( = crab one); C2 = second juvenile Size of of zoea of of Size

Cryptolithodes sitchensis

Z5 = 3.4 and Mc- 2000a) G = 2.2 Laughlin (Crain and 2000a) McLaughlin 2000a)

stages

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal

Number of Number Eclosion date Eclosion

Phyllolithodes papillosus

Oedignathus inermis

April 4-5 Z, 1 G Z1 = 1.1-1.5, PZ (brief) C1 = 2.4 Z1 drawings (Haynes (Crain and Z1 = 5.1 Z1 = 6-8 C2 = 2.8 (Haynes 1993), McLaughlin -6.0 TL Z2 = 5-10 C1 = 32- 1993), ZI-4, G, March 2000a) (Haynes Z3 = 5-9 40 days C1 and C2 (Barkley 1993) Z4 = 6-14 C2 = un- (Crain and Sound, B.C., Z1 = 2.6 Z5 = unknown McLaughlin Canada, Z2 = 3.0 known duration 2000a) Zaklan, Z3 = 3.3 G = 24-29 (Crain and pers. obs.) Z4 = 3.5 (Crain McLaughlin

(mm)

Egg diameter Egg

(eggs/clutch)

Brood size Brood

cycle

Reproductive

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

aldwyn 1970)

CL = 113 (Macpherson 1988c)

CL = 133 (Macpherson 1988c)

CL = 130, CW = 104 Y (Dawson and

turity of males of turity productive ma- productive

(courtesy of Greig Jensen, C. Nyblade, and Hokkaido University), and size (mm) at re- at (mm) size Age (years) and (years) Age

able 5. (Continued.) Life history traits of the family Lithodidae. Species Measurement of carapace length (CL) unless otherwise noted.

Lopholithodes mandtii

Neolithodes agassizii

N. asperrimus

N. brodiei

Crabs in Cold Water Regions: Biology, Management, and Economics 805

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile,

Size of first of Size

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

(Z) glaucothoe (Z) Size of of zoea of of Size

Z1 = 1.36 Yoshida foregut

Z3 = 1.77 and Kittaka G = 1.69 1997) (Nakanishi 1981)

stages

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal

Number of Number

Eclosion date Eclosion

(mm)

Egg diameter Egg

(eggs/clutch) Brood size Brood

8,000-79,000 March/ PZ, 3 Z, 1 G Z1 = 1.41 Z1-G = 35- C1 = 1.78 Z1 and G

3,833 1.62-1.96 pers. obs.) pers. obs.) (K. Rypien, (K. Rypien, U. of Alberta U. of Alberta

cycle Reproductive

oshida 1999b)

Molt, mate and

in June (Sasaki and Z2 = 1.69 1999) (Abrunhosa Y

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 118 (Macpherson 1988c)

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 96.4, CL = 94.5, age 6 (Abe age 6 egg release (Sato and April (Kurata Z2 = 1.49 70 (Tº de- (Kurata (Kurata 1956), and Koike (Abe and in June and Abe 1941) (Kurata 1956) Z3 = 1.68 pendent; 1956) C2 (Sasaki 1982), Koike 1982) July (Sato and 1956) G = 1.60 Nakanishi C2 = 2.28 and Yoshida CL = 70 Abe 1941), (Kurata 1981) (Sasaki 1999), mouth- (Abe 1992)settle larvae 1956) andand parts able 5. (Continued.) Species

californiensis

N. grimaldii

Paralithodes brevipes

P. 806 Zaklan — Review of Lithodidae

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile, Size of first of Size

Acantholithodes hispidus

pendent, (Stevens 1990) C1 CL = 2.18 (Donaldson et al. 1992)

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

(Z) glaucothoe (Z) Size of of zoea of of Size juvenile instar ( = crab one); C2 = second juvenile

Cryptolithodes sitchensis

stages .

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal

Number of Number Eclosion date Eclosion

Phyllolithodes papillosus

Oedignathus inermis

(mm)

Egg diameter Egg

(eggs/clutch)

Brood size Brood

cycle

Reproductive

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 102 (Somerton 1980), CL = 76-105 (Otto et al. 1980), (Paul 1992) CL = 76-88

turity of males of turity productive ma- productive

(courtesy of Greig Jensen, C. Nyblade, and Hokkaido University), and size (mm) at re- at (mm) size Age (years) and (years) Age

Area Area 11-13 month 70,000- 0.71-0.82 March to 4 Z, 1 G Z1 = 1.18 Z1 = 14 C1 CL Egg, Z1-4 and dependent: dependent: cycle, annual 270,000 (Marukawa May (Marukawa Z2 = 1.38 Z2 = 8 = 2.5, G (Marukawa age 8 (Loher CW = 85-100 spring mi- (Marukawa 1930), (Marukawa 1930) Z3 = 1.45 (Marukawa C1 CW = 1930, Sato et al. 2001), (Marukawa gration and 1930), 0.88-1.03 1930) Z4 = 1.53 1930), 2.0 and Tanaka chelae 1930), synchronized 15,330- (Matsuura Z4 = 1.53 Z 1-4 = (Marukawa 1949, Kurata allometry CL = 83-95 spawn 214,410 and G = 1.8 47-84 1930) 1964), egg CL = 103 (Powell and (Marukawa (Sato 1958) Takeshita G = 1.5 CW (Sato C1 CL = (Nakanishi (Somerton 1965b), 1933), mate 1985) (Sato and 1958) 2.50, 1987), mouth 1980), Nickerson February to Tanaka Z1 = 14 C1 CW = and foregut CL = 120, 86-119 April shallow 1949), Z2 = 14 1.33 (Abrunhosa presence of (Powell et al. water (Powell G = 2.0 (Shirley (Sato and and Kittaka sperma- 1973b), age 5 et al. 1973b), G = 1.7 CW and Tanaka 1997) tophores at (McCaughran brood for (Marukawa Shirley 1949), CL = 70-99 and Powell 300 days 1930) 1988) Tº de- (Alaska, Paul 1977), chelae (Nakanishi et al. 1991) allometry 1987) able 5. (Continued.) Life history traits of the family Lithodidae. Species

cam-

Measurement of carapace length (CL) unless otherwise noted.

P. tschaticus

Crabs in Cold Water Regions: Biology, Management, and Economics 807

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe Zoeal (Z), glau- (Z), Zoeal

and Kittaka 1997)

(mm)

juvenile, C1 juvenile,

Size of first of Size

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

(Z) glaucothoe (Z)

Size of of zoea of of Size

stages

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal

Number of Number

Eclosion date Eclosion (mm)

0.98 March to 4 Z, 1 G Z1 = 1.2, 3.2 Z1 = 12.4 2.6 Z1 (Marukawa Egg diameter Egg ¥

1.0 Armstrong (CL, TL) G = 12.8 man 1968),

MacIntosh

(eggs/clutch)

Brood size Brood

cycle Reproductive

(Sasakawa (Sasakawa (Sasakawa (Jensen Z3 = 1.6, 3.5 Z3 = 12.5 1968) 1964), Z1-4

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 38 (Macpherson 1988c)

CL = 68, CW = 68 (Macpherson 1988c)

CL = 48, 1988c) CW = 51 (Macpherson

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

Area Area 19 month 3,000- 1.18 dependent: dependent: cycle 160,000 Asia May (Sato 1958) Z2 = 1.3, 3.4 Z2 = 12.3 (Hoffman 1930, Kurata sperm CL = 80.6-96.3 CL > 70 (Alaska, 1975a), bien- 1975b) 1975b), and Z4 = 2.0, 4.5 Z4 = 14.3 and G (Hoff- (Sasakawa Somerton and nial mating 1.2 1971), MacIntosh and broods Alaska 1989) G = 1.8 CL (Hoffman mouth and CL = 77-108 1983), for 14-15 (Somerton (Hoffman 1968) foregut (Alaska, CL = 101-105 months and 1968) (Ambrunhosa Somerton (Otto et al. (Somerton and and Mac- 1980) MacIntosh 1985) Intosh 1983), 1985), sperma- 12 months tophores in (Jensen and CL = 50-69 Armstrong (Alaska, Paul 1989) et al. 1991)

CL = 38.5, CL = 33.0, CW = 35.0 CW = 32.0 (Pohle 1992b) (Pohle 1992b) able 5. (Continued.) Species

platypus

anamerae

bouvieri

cristulata

Paralomis africana

P. 808 Zaklan — Review of Lithodidae

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile, Size of first of Size

Acantholithodes hispidus

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

(Z) glaucothoe (Z) Size of of zoea of of Size juvenile instar ( = crab one); C2 = second juvenile

Cryptolithodes sitchensis

stages .

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal

Number of Number Eclosion date Eclosion

Phyllolithodes papillosus

Oedignathus inermis

(mm)

Egg diameter Egg

(eggs/clutch) Brood size Brood

eggs and Vinuesa after Guzman G = 2.0 4.8 Vinuesa Guzman (Hoggarth 1993) mating), 1981) G = 2.0 (Campo- 1996) 1981)

(Lovrich and Vinuesa (Campo- Guzman

cycle

Reproductive

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 58, CW = 57 (Macpherson 1988c)

CL = 64 (Macpherson 1988c)

turity of males of turity productive ma- productive

(courtesy of Greig Jensen, C. Nyblade, and Hokkaido University), and size (mm) at re- at (mm) size Age (years) and (years) Age

inuesa 1993, maturity with a 10 1993) Guzman

Gonadal Gonadal Biennial, 50% of 2.3 (Vinuesa June- 1-2 Z, 1 G Z1 = 2.1 Z1 = 5.1- C1 = 3 CL Z1-2 and G maturity maturity mate and adults do 1987), 1.9 August (Campo- Z1 = 5.9- 5.7 (Lovrich (Campo- CL = 50.2 CL = 60.6, molt Oct-Jan. not carry (Lovrich (2 years donico and 6.0 TL Z2 = 4.2- and donico and or age 10, morphometric asynchronous morpho- maturity) development, metric CL = 66.5 eggs extruded 1993), (Lovrich CW G = donico maturity (Lovrich and Oct-Nov, 800-10,000 and 4.5 TL and CL = 57 1993), Vinuesa embryogenesis (Lovrich and functional 18-22 months Vinuesa 1993) donico and 1981) V 1995, 1999), CL = 46 month dia- 1981) functional (Hoggarth pause (Lovrich maturity 1993 and Vinuesa 1993) CL = 52 1993, 1996) (Hoggarth able 5. (Continued.) Life history traits of the family Lithodidae. Species

erinacea

formosa

granulosa

Measurement of carapace length (CL) unless otherwise noted.

Crabs in Cold Water Regions: Biology, Management, and Economics 809

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile,

Size of first of Size

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

(Z) glaucothoe (Z) Size of of zoea of of Size

aishaku 1994)

T 1994)

stages

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal Number of Number

anagisawa Yanagisawa 1985

2 Z and G Z1 = 2.5- PZ and Z1 (Hayashi 2.9 (Hayashi (Hayashi and and and Yanagisawa Y 1985) 1985) Eclosion date Eclosion

aishaku Taishaku (Konishi and Taishaku

April 2 Z and 1 G Z1 = 3.4 Z1 and Z1 and Z2 (Konishi (Konishi Z2 = 3.6 Z2 = 17 and G and and G = 2.5 (Konishi (Konishi and T 1994) 1994) and Taishaku 1994)

(mm)

Egg diameter Egg

(eggs/clutch)

Brood size Brood

cycle

Reproductive

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 98, CW = 98 (Macpherson 1988c)

CL = 49, CW = 48 (Macpherson 1990)

CL = 93, R = 10, CW = 109 (Haig 1974)

CL+R = 56.8, CW = 55.5 1990) (Takeda and (Takeda Hashimoto

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size

Age (years) and (years) Age able 5. (Continued.) Species

grossmani

haigae

hystrix

inca

japonica

jamsteci

P. 810 Zaklan — Review of Lithodidae

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile, Size of first of Size

Acantholithodes hispidus

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

(Z) glaucothoe (Z) Size of of zoea of of Size juvenile instar ( = crab one); C2 = second juvenile

Cryptolithodes sitchensis

stages .

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal

Number of Number Eclosion date Eclosion

Phyllolithodes papillosus

Oedignathus inermis

(mm) Egg diameter Egg

2 (Faxon 1893)

(eggs/clutch)

Brood size Brood

cycle

Reproductive

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL+R = 63, CW = 55 (Takeda 1974) (Takeda

CL = 89.5, CW = 88.3 (Wilson 1990) (Wilson

CL = 96, CW = 96 (Macpherson 1988c)

CL = 59, CW = 63 (Macpherson 1992) (Eldredge CL = 62, 2.5 (Eldredge CW = 57 1976)

turity of males of turity productive ma- productive

(courtesy of Greig Jensen, C. Nyblade, and Hokkaido University), and size (mm) at re- at (mm) size

Age (years) and (years) Age able 5. (Continued.) Life history traits of the family Lithodidae. Species

longipes

otsuae

pectinata

phrixa

seagranti

Measurement of carapace length (CL) unless otherwise noted.

P. medipacifica

P. R = rostrum; CL = carapace length; CW = carapace width; TL = total length; G = glaucothoe; Z = zoea; PZ = pre-zoea; C1 = first instar ( = crab two). McLaughlin and Lemaitre (2000) report the possession of glaucothoeal and first crab stages of Hapalogaster dentata a

Crabs in Cold Water Regions: Biology, Management, and Economics 811

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile,

Size of first of Size

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

(Z) glaucothoe (Z)

Size of of zoea of of Size

stages

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal

Number of Number

Eclosion date Eclosion

(mm)

Egg diameter Egg

(eggs/clutch) Brood size Brood

2,000- 2.0 (Otto

cycle

Reproductive

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CW = 72 (Macpherson 1990)

CL = 47, CW = 45 (Macpherson 1988c)

CL = 49, CW = 50 (Macpherson 1988a)

CL = 41, CW = 47 (Macpherson 1988b) (Sakai 1987)

CL = 99

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 74.8 CL = 61.7 Asynchronous South (Otto and release date 14,000 1993) Georgia, MacIntosh (Otto and (Otto 1993) CL = 66.4 1996) Macintosh Shag Rocks 1996) Antarctic (Otto and MacIntosh 1996)

sp. CL = 71,

able 5. (Continued.)

spectabilis

stella

tuberipes

verrilli Species

P. spinosissima

P. 812 Zaklan — Review of Lithodidae

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile, Size of first of Size

Acantholithodes hispidus

(days)

cothoe stages cothoe

zoeal and glau- and zoeal

Duration of Duration

(G) (mm) (G)

(Z) glaucothoe (Z) Size of of zoea of of Size juvenile instar ( = crab one); C2 = second juvenile

Cryptolithodes sitchensis

stages .

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal Number of Number

1964) Eclosion date Eclosion

Phyllolithodes papillosus

Oedignathus inermis

(mm)

Egg diameter Egg

(eggs/clutch)

Brood size Brood

cycle

Reproductive

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

CL = 44.87, 1,959-8,705 1.07-1.18 March Z1 = 1.29 CL Drawing of Z1 R = 6.35, (K. Rypien, (Zaklan, (Haynes (Haynes and Z2 CW = 39.18 U. of pers. obs.) 1984) 1984) (Haynes 1984) (Zaklan, Alberta, pers. obs.) pers. obs.)

CL = 24 (Macpherson 1988c)

CL = 24.01, R = 5.9, CW = 32.81 (Zaklan, pers. obs.)

CL = 24.01, 4 Z, 1 G Z1-4 and G CW = 5.52 (Kurata (Kurata 1964) (Zaklan, pers. obs.)

CL = 14.71,obs.) 1,162-2,265 0.71-0.84 CW = 12.62 (Zaklan, (Zaklan, (Zaklan, pers. pers. obs.) pers. obs.)

turity of males of turity productive ma- productive

(courtesy of Greig Jensen, C. Nyblade, and Hokkaido University), and size (mm) at re- at (mm) size

Age (years) and (years) Age able 5. (Continued.) Life history traits of the family Lithodidae. Species

Measurement of carapace length (CL) unless otherwise noted.

Rhinolithodes wosnessenskii

Sculptolithodes derjugini

Subfamily Hapalogastrinae Acantholithodes hispidus

Dermaturus mandtii

Hapalogaster cavicauda R = rostrum; CL = carapace length; CW = carapace width; TL = total length; G = glaucothoe; Z = zoea; PZ = pre-zoea; C1 = first instar ( = crab two). McLaughlin and Lemaitre (2000) report the possession of glaucothoeal and first crab stages of Hapalogaster dentata a

Crabs in Cold Water Regions: Biology, Management, and Economics 813

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile,

Size of first of Size

(days)

cothoe stages cothoe

zoeal and glau- and zoeal Duration of Duration

1961)

(G) (mm) (G)

(Z) glaucothoe (Z) Size of of zoea of of Size

(Konishi

G = 1.98 CL Z4 = 12 1961) G = 1.3, CW G = 16 G = 3.25 TL (Miller (Miller and and Coffin 1961) Coffin

stages

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal Number of Number

1961) Z4 = 5.5 TL Z3 = 12 Coffin

Eclosion date Eclosion

(mm)

Egg diameter Egg

(eggs/clutch)

Brood size Brood

cycle Reproductive

brooding (Goshima (Goshima March (Konishi 1.89 19-23 CL C1 = (Konishi

pers. obs.)

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

age 2 (Goshima

CL = 18, 900-1,800 February- PZ, 4 Z PZ = 3.30 TL PZ = 5-6 C1 = 2.0 Z1-4, G and CW = 16 (Miller and March 1 G Z1 = 3.65 TL hours CL C2 = C1 (Miller and (Miller and Coffin 1961) (Zaklan, (Miller and Z2 = 4.35 TL Z1 = 8.5 2.05 CL Coffin 1961) Coffin 1961) pers. obs.) Coffin Z3 = 4.6 TL Z2 = 11 (Miller and

CL = 10.2, Annual, 208-2,421 1.16-1.19 February- Z1-4 = 30 CW = 10.3 spawn in (Zaklan, (Zaklan, March (Zaklan, (Zaklan, August pers. obs.) pers. obs.) (Zaklan, pers. obs.) pers. obs.) (Zaklan, pers. obs.)

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size Age (years) and (years) Age

Morph CL = 6.5-8 180 day 73-3,051 0.9-1.04 February- 4 Z, 1 G Z1 = 1.73- Z1-4 = C1 = 1.71 Z1-4 and G maturity CL = 5.2, et al. 1995) period, et al. 1995) et al. 1995), release 1986) Z2 = 1.74- Z1-C1>30 1.21 CW 1986) gonadal (Takahashi 1.14-1.19 (Goshima 1.93 (Konishi (Konishi maturity et al. 1985), (Zaklan, et al. 1995) Z3 = 1.98- 1986) 1986) CL = 5.3 or seasonal, pers. obs.) 2.13 age 1, spawn from Z4 = 2.09- functional Oct. to Nov. 2.37 maturity 110 day G = 1.53- CL = 9.5 brooding 1.64 (Goshima period et al. 1995, (Goshima et 1986) 2000) al. 1995)

able 5. (Continued.) Species

H. dentata

H. mertensii

Oedignathus inermis 814 Zaklan — Review of Lithodidae

drawings

juvenile (C) juvenile

cothoe (G), or (G), cothoe

Zoeal (Z), glau- (Z), Zoeal

(mm)

juvenile, C1 juvenile, Size of first of Size

Acantholithodes hispidus

(days)

cothoe stages cothoe

zoeal and glau- and zoeal Duration of Duration

and Mc- Laughlin 2000b)

(G) (mm) (G)

(Z) glaucothoe (Z) Size of of zoea of of Size juvenile instar ( = crab one); C2 = second juvenile

Cryptolithodes sitchensis

Z3 = 4.5 Z3 = 2000b) Z4 = 4.8 10-13 G = 3.1 Z4 = (Crain and 12-15 McLaughlin G = 10-15 2000b) (Crain

. stages

glaucothoe (G) glaucothoe

zoeal (Z) and (Z) zoeal Number of Number

McLaughlin Z1 = 3.1 Z2 = McLaughlin and 2000b) Z2 = 3.7 10-15 2000b) McLaughlin Eclosion date Eclosion

Phyllolithodes papillosus

Oedignathus inermis

March PZ, 4 Z Z1 = 2.12 PZ = C1 = Z1 (Haynes (Haynes 1 G (Haynes 2 min. 2.6 CL 1984) Z1-4 1984) (Crain and 1984) Z1 = 8-12 (Crain and and G (Crain

(mm)

Egg diameter Egg

(eggs/clutch)

Brood size Brood

cycle

Reproductive

turity of males of turity

productive ma- productive

size (mm) at re- at (mm) size

Age (years) and (years) Age

turity of males of turity productive ma- productive

(courtesy of Greig Jensen, C. Nyblade, and Hokkaido University), and size (mm) at re- at (mm) size Age (years) and (years) Age

able 5. (Continued.) Life history traits of the family Lithodidae. Species Measurement of carapace length (CL) unless otherwise noted.

Placetron wosnessenskii

Table 6. Predator and prey relationships of the family Lithodidae.

Adult Larval Species diet diet Predators

Subfamily Lithodinae Cryptolithodes Calcareous algae, Corallina, Calliar- sitchensis thron, and Bossiella (Hart 1982) and other sessile organisms (Jensen 1995)

C. typicus Opportunistic grazer on bryozoans, coralline algae, and other encrusting or sessile organisms (Hart 1982, Jensen 1995)

Lithodes Lecithotrophic aequispinus (Shirley and Zhou 1997)

L. maja Lecithotrophic (Anger 1996)

L. murrayi Benthic opportunist omnivore includes polychaetes, echinoderms, sponges, gastropods, small crustaceans (Arnaud and Do-Chi 1977)

L. panamensis Sperm whales (Physeter catodon Haig 1974)

L. santolla Broad opportunists consisting of Consume plankton mollusks (mainly gastropods), in zoea stage I, crustaceans, and Bryozoa; prey was nonfeeding in zoea size class and season dependent; stage II, and consumption greatest in wintering glaucothoe, thus large crabs; autumn diet of Pseude- facultatively chinus magellanicus (66.7%); winter lecithotrophic diet of Bryozoa (Membranipora (Comoglio and isabellean); spring diet of Crustacea Vinuesa 1991); e.g. Isopoda and Munida spp.; lecithotrophic summer diet of algae, Macrocystis (Oyarzun 1992) pirifera, feed on more mobile organisms than P. granulosa (Comoglio et al. 1990, Comoglio and Amin 1996)

Lopholithodes A deposit feeder, a filterer of chelae- foraminatus dredged sediment and an opportunist consumer of buried prey such as clams (Jensen 1995)

L. mandtii Opportunist feeder, e.g. echinoderms and sea anemones (Hart 1982) mussels in the lab (Zaklan, pers. obs.)

Neolithodes diomedeae Sperm whales (Baez et al. 1986)

Paralithodes Laminaria longissima and Glaucothoe a non- brevipes Corallina pilulifera (Sasaki and feeding stage Kuwahara 1999) (Abrunhosa and Kittaka 1997) 816 Zaklan — Review of Lithodidae

Table 6. (Continued.) Predator and prey relationships of Lithodidae.

Adult Larval Species diet diet Predators

P. Adults are opportunist omnivores See Paul et al. 1989 Egg predator, camtschaticus (see Takeuchi 1959, 1967, Feder for overview, diel Carcinonemertes et al. 1980, Jewett and Feder 1982 feeding patterns regicides (Nem- for overviews) feeding on mollusks exist (Shirley and ertea; Shields et al. (bivalve families Tellinidae and Shirley 1987) newly 1989, Kuris et al. Cardiidae, gastropods family hatched zoeae are 1991). For over- Trochidae), Crustacea (mainly herbivorous and view see Loher et barnacles), fish, annelids, Polychaeta, carnivory increases al. 1998, predators echinoderms and algae (Takeuchi with age (Shirley include Pacific hal 1959, 1967, Tarverdieva 1976, and Shirley 1989b) ibut (Hippoglossus Jewett and Feder 1982), kelp zoea feed and glau- stenolepis), Pacific (Laminaria sp.), Ulva sp., molt cothoe are non- cod (Gadus macro exuvia, sea stars (Evasterias feeding (Abrunhosa cephalus), sable troschellii and Pycnopodia and Kittaka 1997) fish (Anoplopoma helianthoides; Dew 1990). Young phytoplankton prey fimbria), various juveniles forage through sediment include diatoms, flatfish, flounders eating crustaceans, polychaetes, barnacle and crab (Atheresthes spp.), diatoms, tintinnids, foraminiferans, larvae (Bright 1967) sole, herring, algae and bryozoans (Feder et al. zooplankton such salmon and 1980). as Artemia, sculpins yellow Thalassiosira spp., irish lords (Hemi- Skeletonema lepidotus), snailfish costatum,(Liparis sp.), Chaetoceros spp., eelpout (Lycodes), copepod nauplii, skates (Raja spp.) cannibalism (Kurata and arrowtooth 1959, Paul et al. 1989)

P. platypus Use chelae for crushing mollusks As glaucothoe mouth and urchin testes, feed on hard and parts atrophy (Sato soft bottoms where they excavate and Tanaka 1949) large pits (Somerton 1985) they are a nonfeeding stage (Abrunhosa and Kittaka 1997)

Paralomis bouvieri Northern wolffish Anarhichas denticulatus (Pohle 1992b)

P. formosa Toothfish Dissostichus eliginoides (Konforkin and Kozlov 1992)

P. granulosa Algae, Foraminifera, Bryozoa, Lecithotrophic bivalves, gastropods, barnacles, (Campodonico and ascidians and Polychaeta (Comoglio Guzman 1981)

P. spinosissima Toothfish (Konforkin and Kozlov 1992) Crabs in Cold Water Regions: Biology, Management, and Economics 817

Table 6. (Continued.)

Adult Larval Species diet diet Predators

Phyllolithodes In captivity they eat small sea urchins papillosus (Jensen 1995) and mussels (Zaklan, pers. obs.) and sponges in the wild (Jensen 1995)

Subfamily Hapalo- gastrinae Acantho- Found foraging in prawn traps, most lithodes likely unable to catch shrimp under hispidus natural conditions (Jensen 1995)

Dermaturus Consumer of algae or algal detritus mandtii (Jensen 1995)

Hapalogaster Filter feeds and is a broad cavicauda opportunist omnivore (Jensen 1995)

H. mertensii Filter feeds and is a broad Glaucothoe are non- opportunist omnivore (Jensen 1995) feeding (Miller and Coffin 1961)

Oedignathus Omnivorous and a filter feeder, inermis captive specimens consume worms and crustaceans and crushed mussels (Jensen 1995)

Placetron A fast lithodid predator, forcep-like Planktonic omnivore wosnessenskii chelae are used to obtain crevice- (Crain 1999) dwelling prey such as brittle stars, shrimp, amphipods, crabs and brachiopods (Jensen 1995) 818 Zaklan — Review of Lithodidae

Stegopoma

Poecilasma

and Hydroid -

(Abello and Macpherson

Epibionts: cirripede

kaempferi plicatile 1992)

Isopods, hydroids and polychaetes (spirorbs; Arnaud and Do-Chi 1977)

hyperparasite (Pohle 1992a)

Cryptoniscinid isopod

Commensals

and unknowns

cariae, larval

acanthocephalans, parasitic dinoflagellates, viral

of Nosematidae family producing “cottage cheese disease” (Sparks and Morado 1985)

Eggs exposed to cadmium and lead resulted in early eclosion and larval hatching decrease (Amin et al. 1998)

Other

parasites etc.

, red metacer Trematode

C. falklandica

, Melville-Smith

C. furcellus

infects males only

sp. (Love and Shirley

L. tropicalis

sp. (Campodonico and

sp.

Pink snailfish snailfish 1993, Somerton and Donaldson 1998) infection, microsporidian

C. griseldea (as

and Louw 1987)

C. Guzman 1977), (Balbontin et al. 1979)

(Parrish 1972),

callosus

Briarosaccus Careproctus

sp. (Peden and Corbett 1973)

oshihara 1970; Sloan 1984;

Boschma 1970; McMullen and Y Bower and Sloan 1985; Sparks and Morado 1985; Hawkes et al. 1985a, 1986 Abello and Macpherson 1992

Boschma 1970, Somerton 1981

Arnaud and Do-Chi 1977

Boschma 1970

Boschma 1930, 1970

Pohle 1992a

able 7. Parasites and commensals of the family Lithodidae.

Species

Subfamily Lithodinae Lithodes aequispinus

L. couesi

L. ferox

L. murrayi

L. santolla

Lopholithodes C. melanurus foraminatus C.

Neolithodes agassizii

N. grimaldii Crabs in Cold Water Regions: Biology, Management, and Economics 819

Myti-

Ischyrocerus

), copepods and

larvae (Jansen et al. 1998)

lus edulis

Myzotarsa anaxiphilius (Gammaridea: Amphipoda, Cadien and Martin 1999) Amphipods ( commensalis

Commensals

and unknowns

, Gibson

sp., Sparks

), Acanthocephala

, Jansen et al.

cariae, larval

Carcinonemertes

Alaxinus oclairi

Thelohania

Shields et al. 1989, Kuris

ematode metacer

Johanssonia arctica Profilicollis botulus

Sparks and Morado 1985), parasitic dinoflagellates, microsporidian ( gut (Sparks and Morado 1987), et al. 1990), Protozoa, gill ciliates, and Morado 1985) herpesviridae Acanthocephalans (Sparks 1987), flagellates, turbelarians (Promesos- (Sparks and Morado 1985, 1986), tomidae?), Nemertea, Hirudinea Tr infection ( Nemertea (e.g. regicides, ( et al. 1991; ( 1998)

cercariae, microsporidian

Parasitic dinoflagellates, viral

rickettsiae (Johnson 1984)

Other

parasites etc.

(Rass 1950, infection, trematode meta-

C. sinensis

sp. (Anderson and Cailliet sp. (Nakazawa 1915, Hunter

sp.

inogradov 1950)

1974) 1969), V

C. C.

sp., acanthocephalans, viral infection,

oshihara 1970,

callosus

Briarosaccus Careproctus

Thompsonia

1987; Hawkes et al. 1986b, Jansen Johnson et al. 1986 et al. 1998 carcinoma-like growth in hind-

Hawkes et al. 1985a,b, 1986a, Sparks and Morado 1985,

Donald Cadien, Redondo Submarine Canyon, Los Angeles, U.S.A. at 305 Haynes 1969, Boschma 1970, m: 33º 49.23/118º 27.09: McMullen and Y This study, collected by This study, August 6, 1997 Faxon 1895, Boschma and

able 7. (Continued.)

platypus

camtschaticus

P. T

californiensis

Species

Paralithodes P. 820 Zaklan — Review of Lithodidae

and Martin 1999)

- Myzotarsa anaxiphilius (Gammaridea: Amphipoda, Cadien

Commensals

and unknowns

(Otto and

(Isopoda:

akeda and

B. callosus

Cryptoniscinid isopod hyper Microsporidian infection (possibly Pseudione tuberculata family Nosematidae) isopod hyper infested by a sacculinid parasite parasite on (Takeda and Ohta 1979, Sakai 1987) (Takeda Bopyridae; Roccatagliata and Lovrich 1999) MacIntosh 1996)

Sacculinid parasite (T Miyake 1980)

Other

parasites etc.

sp. (Balbontin et al. 1979)

sp.

(Boschma

californiensis

callosus

Briarosaccus Careproctus

above) Pohle 1992b parasite (Pohle 1992b)

This study, collected by D. This study, Faxon 1895, Lutzen 1987 Cadien, Redondo Submarine Only males infected, Canyon (see Chace 1939 Otto and MacIntosh 1996 Lutzen 1985 Boschma 1970

Zaklan, pers. obs. (Barkley Sound, B.C., 20 m)

1970, Walossek et al. 1996) 1970, Walossek

able 7. (Continued.) Parasites and commensals of the family Lithodidae.

rathbuni

cubensis spinosissima granulosa truncatispinosa

bouvieri Species Paralomis cristata P. Paralomis aspera Paralomis P. P. P. P. P.

Phyllolithodes papillosus

Subfamily Hapalogastrinae Hapalogaster Briarosaccus tenellus

mertensii Crabs in Cold Water Regions: Biology, Management, and Economics 821

inuesa

tment of Fish and Game’

references

Hiramoto 1985, Otto et al. 1990, Jewett et al. 1985,

Orensanz et al. 1998

Macpherson 1988c, V

et al. 1996 Arnaud et al. 1976

Suggested

L. santolla

camtschat-

brevipes

and overlaps

and

the fishery is unknown

Age of recruitment into

icus

(592-850 m), generally as bycatch to aequispinus

Morphology similar to A small fishery, dueA small fishery, Somerton 1981 L. santolla

in distribution thus likely Exploratory fishery to be landed as in places, fishery opening soon in Argentina

Comments

efer to fishery internet sites including Alaska Depar

) traps of to deep water habitat

. Shirley,

camtschaticus

2.5 m that are

0.8

method

Anoplopoma fimbria

longlines in Aleutians, Bristol

Mesh covered pots are set on

U.S.A. Pot limits vary with area, and depend on guideline harvest levels, vessel size and biomass traps (see below) arranged at estimates. In southeastern 91.5 m intervals with surface Alaska, pot limits vary between floats connected to both ends 20 and 250 pots (T are used. pers. comm.). covered with 8.9 cm webbing, or rectangular

0.8 m

two kinds of PVC pots (“Alaskan” king crab pots and regular

Caught using beam trawl and

lobster pots).

eflect changes in the population, please r

of males

dependent, 5.0

Area and year

to 7.0 inches Bay and Bering Sea, Alaska,

5.5 inches Both rectangular sablefish

e modified year to year to r

Central Japan, along Aleutian Islands and

Sea, Alaska, U.S.A. to southern B.C., Canada

(S.W. Indian Ocean) (S.W.

Crozet Islands

Area Minimum size Fishing

able 8. Summary information concerning fisheries of the family Lithodidae.

As harvest guidelines ar

Species harvested

Lithodes aequispinus (brown or continental slope of

golden king) southeastern Bering

L. confundens L. couesi (deep sea or (140 mm) ( scarlet king crab)

L. murrayi

commercial shellfish regulations for up-to-date information such as opening dates, minimum harvest sizes, and catch limits. 822 Zaklan — Review of Lithodidae

references

yngaard and Iorio 1996;

inuesa 1999

W Campodonico 1983; Boschi et al. 1984; Bertuche et al. 1985; Lovrich and Vinuesa 1996, Lovrich and Vinuesa 1999; Vinuesa et al. 1996; 1999; Vinuesa

Abe 1992

Suggested

ry small fishery Baez et al. 1986, Lovrich and

camtschaticus

fisheries in the Southern Hemisphere until the collapse in 1993. Now open One of the largest In theperiodically. Lovrich 1997 Golfo San Jorge it is a trawling fishery with high population impact as there is no sex or size selectivity nor returns of sublegal crabs.

Increasing in size with the decline of Ve

Comments

Merluccius

in the Atlantic. Pots

)

method

en pots are longlined and en pots are

Argentinean hake ( (Chilean design) or the more efficient conical traps (Japanese design), or illegally with tangle nets. Often a bycatch of the

hubbsii are truncated cones (1.3 m high) of three iron hoops united by eight crossbars of thinner iron rods covered with mesh. Pots have a base diameter of 1.5-1.8 m and opening entrance of 0.6-0.8 m with a circular plastic escape guard. T each separated by 20 m.

of males

CL = 120 Caught legally with spherical

Argentina (Beagle of Magellan (Chile)

city of Comodoro Rivadavia) and Strait

Mostly a Japanese fishery, in the Sea of fishery, Puerto Mont, Argentina

Japan

Area Minimum size Fishing

able 8. (Continued.) Summary information concerning fisheries of the family Lithodidae.

Species harvested

L. santolla (centolla, Channel and Golfo San formerly Jorge at 46ºS near the L. antarcticus)

Paralithodes brevipes Neolithodes diomedeae

(Hanasaki crab) Crabs in Cold Water Regions: Biology, Management, and Economics 823

inuesa

tment of Fish and Game’

references

Loher et al. 1998; Kruse et al. 2000; Zheng and Kruse 2000 Otto 1990; Zheng et al. Marukawa 1930, 1933; Otto 1990; Abe 1992; Zheng et al. 1997a; Orensanz et al. 1998;

1997b, 1998; Orensanz et al.

1998

et al. 1996; Lovrich 1997; Lovrich and Vinuesa 1993, Lovrich and Vinuesa

Campodonico 1983; V

Suggested

L. santolla

are higher are 1995, 1999

, and since the

granulosa

spring, molting and mating periods. estimates) are attained (between 7-10 days). The fishery occurs in the fall months until quotas

Exploratory fishery, Otto and MacIntosh 1996 A co-fishery with July-November; fishery opening soon in early 1990s landings of Argentina than those of

fishery opening soon in Annual catch of 3,000 t.

Exploratory fishery, Otto and MacIntosh 1996

Comments

efer to fishery internet sites including Alaska Depar

0.9 covered (based on population

1.8

L. santolla santolla

method

by polypropylene mesh with two

legal commercial fishing gear and

eflect changes in the population, please r

of males CW = 165 mm Age of recruitment into years of age by 89 cm. There is no fishing during

U.S.A.), 8-9 side tunnel openings of 18.5 (6.5 inches) the fishery is unknown CW = 5.5- In Alaska, crab pots are the only 8 inches (varies with harvest measure 1.8

Pribilof Islands;

(5.5 inches) St. Matthew Island CW = 90 mm Caught using similar nets and CL = 82 mm traps as

South Georgia January and November;

CW = 94 mm at Shag Rocks Argentina

e modified year to year to r

Pribilof and St. Matthew islands in Aleutians Northern B.C. Canada,

(Chukchi Sea to

SE Alaska) CW = 140 mm

South Georgia area Antarctic Ocean Magellan (Chile)

Argentina (Beagle Channel) and strait of

Ocean CW = 84 mm at

Chile/south Georgia area of the Southern

Area Minimum size Fishing

able 8. (Continued.)

platypus

camtschaticus

granulosa

spinosissima

As harvest guidelines ar

Species harvested red king crab) P. (blue king crab) P. (Alaskan, U.S.A., Japan, and Russian, Russia, beginning in Japanese, Norway district in Alaska,

Paralomis formosa

P. (false centolla)

a commercial shellfish regulations for up-to-date information such as opening dates, minimum harvest sizes, and catch limits. 824 Zaklan — Review of Lithodidae

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Takeda, M., and S. Ohta. 1979. A new species of the Lithodidae (Crustacea, Anomu- ra) from Suruga Bay, central Japan. Bull. Natl. Sci. Mus. Ser. A (Tokyo) 5:195- 200. Takeshita, K., H. Fujita, and S. Kawasaki. 1978. Carapace moire photography in crabs. Bull. Far Seas Fish. Res. Lab. (Shimizu) 16:115-119. Takeuchi, I. 1959. Food of king crab (Paralithodes camtschatica) off the west coast of Kamchatka in 1958 (Nishi Kamchatka Okiai ni okeru Tarabagani no Shiryo). Bull. Hokkaido Reg. Fish. Res. Lab. 20:67-75. (Translated 1968, by Translation Bureau for Fisheries Research Board of Canada No. 1193.) Takeuchi, I. 1967. Food of king crab (Paralithodes camtschatica) off the west coast of Kamchatka Peninsula, 1958-1964. Bull. Jpn. Sea Reg. Fish Res. Lab. 33:32- 44. (Translated, 1968, by Translation Bureau for Fisheries Research Board of Canada No. 1194.) Tarverdieva, M.I. 1976. Feeding of the Kamchatka king crab Paralithodes camtschatica and Tanner crabs Chionoecetes bairdi and Chionoecetes opilio in the southeastern part of the Bering Sea. Biol. Morya 2:34-39. Tilesius, W.C. 1815. De Cancris Camtschaticis, Oniscis, Entomostracis et Cancellis marinis microscopicis noctilucentibus, Cum tabulis IV: Aenaeis et appendice adnexo de Acaris et Ricinis Camtschaticis. Auctore Tilesio. Conventui exhibuit die 3 Februarii 1813. Memoires de l’Academie Imperiale de Sciences de St. Petersbourg 5:331-405. Tracey, G.A., E. Saade, B. Stevens, P. Selvitelli, and J. Scott. 1998. Laser line scan survey of crab habitats in Alaskan waters. J. Shellfish Res. 17:1483-1486. Tudge, C., B.G.M. Jamieson, L. Sandberg, and C. Erseus. 1998. Ultrastructure of the mature spermatozoon of the king crab Lithodes maja (Lithodidae, Anomura, Decapoda): Further confirmation of a lithodid-pagurid relationship. Invertebr. Biol. 117:57-66. Vinogradov, K.A. 1950. K biologii Tikhookeanskogo pinagora v Kamchatskikh vodakh (Biology of the Pacific lumpfishes in Kamchatka waters). Priroda (Leningrad) 39:69-70. Vinogradov, L.G. 1968. The crab population in Kamchatka. Priroda 7:43-50. Vinuesa, J.H. 1982. Biologia de la reproduccion y el desarrollo embrionario y larval de la centolla, Lithodes antarcticus Jacquinot, en el Canal Beagle, Tierra del Fuego. Ph.D. thesis, University of Bueno Aires. Vinuesa, J.H. 1984. Sistema reproductor, ciclo y madurez gonadal de la centolla (Lithodes antarcticus), del Canal Beagle. In: Estudio Biológico pesquero de la centolla (Lithodes antarcticus) del Canal Beagle, Tierra del Fuego, Argentina. Segunda parte. Contrib. INIDEP (Instituto Nacional de Investigación y Desar- rollo Pesquero), Mar del Plata 441:73-95. Vinuesa, J.H. 1987. Embryonic development of Lithodes antarcticus Jacquinot. De- velopmental stages, growth and mortality. Physis Secc. A 45(108):21-29. Vinuesa, J.H. 1991. Biologia y pesqueria de la centolla (Lithodes santolla). Atlantica, Rio Grande 13:223-244. 844 Zaklan — Review of Lithodidae

Vinuesa, J.H., L. Ferrari, and R.J. Lombardo. 1985. Effect of temperature and salinity on larval development of southern king crab (Lithodes antarcticus). Mar. Biol. 85:83-87. Vinuesa, J.H., L. Guzman, and R. Gonzalez. 1996. Overview of southern king crab and false king crab fisheries in the Magellanic region. In: High latitude crabs: Biology, management, and economics. University of Alaska Sea Grant, AK-SG- 96-02, Fairbanks, pp. 3-11. Vinuesa, J.H., G.A. Lovrich, and L.I. Comoglio. 1991. Madurez sexual y crecimiento de la hembra de centolla Lithodes santolla (Molina, 1782) en el Canal Beagle. Biota, Osorno, Chile 7(1-2):7-13. Walossek, D., J.T. Hoeg, and T.C. Shirley. 1996. Larval development of the rhizocephalan cirripede Briarosaccus tenellus (Maxillopoda: Thecostraca) reared in the laboratory: A scanning electron microscopy study. Hydrobiologia 328:9- 47. Wada, S., M. Ashidate, and S. Goshima. 1997. Observations on the reproductive behavior of the spiny king crab Paralithodes brevipes (Anomura: Lithodidae). Crustac. Res. 26:56-61. Wicksten, M. K. 1987. Range extensions of offshore decapod crustaceans from Cal- ifornia and western Mexico. Calif. Fish Game 73:54-56. Wicksten, M.K. 1989. Ranges of offshore decapod crustaceans in the eastern Pacific Ocean. Trans. San Diego Soc. Nat. Hist. 21:291-316. Williams, A. B. 1984. Shrimps, lobsters and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washing- ton, D.C. 550 pp. Williams, A.B., C.R. Smith, and A.R. Baco. 2000. New species of Paralomis (Decapo- da, Anomura, Lithodidae) from a sunken whale carcass in the San Clemente Basin off southern California. J. Crustac. Biol. 2:281-285. Wilson, R. 1990. Paralomis otsuae, a new species of Decapoda Anomura from deep water off the Chilean coast. Crustaceana 58:130-135. Wu, S-H., T-Y. Chan, and H-P. Yu. 1998. First record of the king crab Lithodes turritus Ortmann, 1892 (Decapoda, Lithodidae) in Taiwanese waters. Crustaceana 71:818-823. Wyngaard, J.G., and M.I. Iorio. 1996. Status of the southern king crab (Lithodes santolla) fishery of the Beagle Channel, Argentina. In: High latitude crabs: Biol- ogy, management, and economics. University of Alaska Sea Grant, AK-SG-96- 02, Fairbanks, pp. 25-39. Yaldwyn, J.C., and E.W. Dawson. 1970. The stone crab Lithodes murrayi Henderson: the first New Zealand record. Records of the Dominion Museum 6:275-284. Yumao, T., W. Fuzhen, and L. Zhicheng. 1984. A new species of Lithodidae (Crusta- cea, Anomura) from East China Sea. Zool. Res. 5:329-332. Zaklan, S.D. 2000. A case of reversed asymmetry in Lithodes maja (Linnaeus, 1758) (Decapoda, Anomura, Lithodidae). Crustaceana 78:1019-1022. Crabs in Cold Water Regions: Biology, Management, and Economics 845

Zaklan, S.D., 2001. Evolution of the family Lithodidae (Crustacea, Anomura, Paguroi- dea). Ph.D. thesis, University of Alberta, Edmonton, Alberta. 280 pp. Zheng, J., and G.H. Kruse. 2000. Recruitment patterns of Alaskan crabs in relation to decadal shifts in climate and physical oceanography. J. Mar. Sci. 57:438- 451. Zheng, J., M.C. Murphy, and G.H. Kruse. 1995. A length-based population model and stock-recruitment relationships for red king crab, Paralithodes camtschaticus, in Bristol Bay, Alaska. Can. J. Fish. Aquat. Sci. 1229-1246. Zheng, J., M.C. Murphy, and G.H. Kruse. 1997a. Analysis of harvest strategies for red king crab, Paralithodes camtschaticus, in Bristol Bay, Alaska. Can. J. Fish. Aquat. Sci. 54:1121-1134. Zheng, J., M.C. Murphy, and G.H. Kruse. 1997b. Application of a catch-survey analysis to blue king crab stocks near Pribilof and St. Matthew Islands. Alaska Fish. Res. Bull. 4:62-74. Zheng, J., M.C. Murphy, and G.H. Kruse. 1997c. Alternative rebuilding strategies for the red king crab Paralithodes camtschaticus fishery in Bristol Bay, Alaska. J. Shellfish Res. 16:205-217. Zheng, J., M.C. Murphy, and G.H. Kruse. 1998. Abundance estimation of St. Mat- thew Island blue king crabs using survey and commercial catch and effort data. In: F. Funk, T.J. Quinn II, J. Heifetz, J.N. Ianelli, J.E. Powers, J.F. Schweigert, P.J. Sullivan, and C.-I. Zhang (eds.), Fishery stock assessment models. Univer- sity of Alaska Sea Grant, AK-SG-98-01, Fairbanks, p. 1037. Zhou, S., and T.C. Shirley. 1998. A submersible study of red king crab and Tanner crab distribution by habitat and depth. J. Shellfish Res. 17:1477-1479. Zhou, S., T.C. Shirley, and G.H. Kruse. 1998. Feeding and growth of the red king crab Paralithodes camtschaticus under laboratory conditions. J. Crustac. Biol. 18:337-345.

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Participants

Alex Andrews Forrest Blau USGS Alaska Biological Science Ctr. 10783 Birch Circle Glacier Bay Field Station Kodiak AK 99615 P.O. Box 240009 907-486-1853 Douglas AK 99824 [email protected] 907-364-1568 [email protected] Bodil Bluhm Alfred-Wegener-Institute Claire Armistead Columbusstrasse National Marine Fisheries Service Bremerhaven 27515 301 Research Court GERMANY Kodiak AK 99615 [email protected] 907-481-1730 [email protected] Jennifer Boldt University of Alaska Fairbanks David Barnard JC-SFOS Alaska Dept. of Fish & Game 11120 Glacier Hwy. 211 Mission Rd. Juneau AK 99801 Kodiak AK 99615 907-465-6441 907-486-1875 [email protected] [email protected] Linda Brannian Bill Bechtol Alaska Dept. of Fish & Game Alaska Dept. of Fish & Game 333 Raspberry Rd. 3298 Douglas St. Anchorage AK 99518-1599 Homer AK 99603-7942 907-267-2118 907-235-8191 [email protected] [email protected] Liz Brown Fran Bennis Alaska Dept. of Environmental Alaska Oceans Network Conservation 441 W 5th Ave. #402 P.O. Box 266 Anchorage AK 99501 Unalaska AK 99685 907-929-3553 [email protected] [email protected] AnnDorte Burmeister Gretchen Bishop Greenland Institute of Natural Resources Alaska Dept. of Fish & Game Box 570 P.O. Box 240020 Nuuk DK-3900 Douglas AK 99824-0020 GREENLAND 907-465-4269 [email protected] [email protected] Ryan Burt Jim Blackburn Alaska Dept. of Fish & Game Alaska Dept. of Fish & Game P.O. Box 920587 211 Mission Rd. Dutch Harbor AK 99692 Kodiak AK 99615 907-581-1239 907-486-1863 [email protected] [email protected] 848 Participants

Susan Byersdorfer Fritz Funk Alaska Dept. of Fish & Game Alaska Dept. of Fish & Game 211 Mission Rd. P.O. Box 25526 Kodiak AK 99615 Juneau AK 99802 [email protected] 907-465-6113 [email protected] John Clark Alaska Dept. of Fish & Game Tamara Gage P.O. Box 240020 Port Gamble S’Klallam Tribe Douglas AK 99824-0020 Natural Resources Dept. 907-465-4245 31912 Little Boston Rd, NE [email protected] Kingston WA 98346 [email protected] James Coe NOAA/NMFS Caleb Gardner Alaska Fisheries Science Center Tasmanian Aquaculture & Fisheries 7600 Sand Point Way NE Institute Seattle WA 98115 P.O. Box 252-49 206-526-4000 Hobart TAS 7004 [email protected] AUSTRALIA +61 3 6277 7233 Peter Cummiskey [email protected] National Marine Fisheries Service P.O. Box 1638 Lee Ann Gardner Kodiak AK 99615 RWJ Consulting 907-481-1720 P.O. Box 672302 [email protected] Chugiak AK 99567-2302 907-688-1400 Earl Dawe [email protected] Dept. of Fisheries & Oceans P.O. Box 5667 Kirsten Gravel St. John’s NFLD Institut Maurice-Lamontagne A1C 5X1 CANADA Fisheries & Oceans Canada [email protected] 850, route de la Mer CP 1000 Mont-Joli Braxton Dew Quebec G5H 3Z4 National Marine Fisheries Service CANADA 7600 Sand Point Way NE BIN C15700 David Hankin Seattle WA 98115-0079 Humboldt State University 206-526-4132 Dept. of Fisheries [email protected] Arcata CA 95521 707-826-3683 Karen DiBari [email protected] Alaska Marine Conservation Council P.O. Box 101145 Gretchen Harrington Anchorage AK 99510 National Marine Fisheries Service 907-227-5357 P.O. Box 21668 [email protected] Juneau AK 99802 907-780-6741 Wes Ford [email protected] Dept. of Primary Industries, Water & Environment Randy Hatch GPO Box 44A Point No Point Treaty Council Hobart TAS 7001 7999 NE Salish Lane AUSTRALIA Kingston WA 98346 [email protected] 360-297-3422 [email protected] Crabs in Cold Water Regions: Biology, Management, and Economics 849

Tracy Hay Steve Jewett NT DPIF University of Alaska Fairbanks P.O. Box 990 SFOS Darwin NT 0801 P.O. Box 757220 AUSTRALIA Fairbanks AK 99775-7220 [email protected] 907-474-7841 [email protected] Nauja Heiselberg Greenland Institute of Natural Resources Christopher Jones P.O. Box 570 NMFS Nuuk DK-3900 Southwest Fisheries Science Center GREENLAND 8604 La Jolla Shores, Drive [email protected] La Jolla CA 92937 619-546-5605 Sandy Hinkley [email protected] Alaska Dept. of Fish & Game P.O. Box 240020 Knut Jørstad Douglas AK 99824 Institute of Marine Research [email protected] Dept. of Aquaculture Postbox 1970 Nordnes Ann Merete Hjelset Bergen N-5817 Norwegian Institute of Fisheries & NORWAY Aquaculture 475-523-6347 N-9291 Tromso [email protected] NORWAY [email protected] Jiro Kittaka Nemuro City Fisheries Res. Inst. Sung Yun Hong 168 Onnemoto Pukyong National University Nemuro Dept. of Marine Biology Hokkaido 087-0166 599-1 Daeyeon-dong, Nam-gu JAPAN Pusan 608-737 [email protected] KOREA [email protected] Tom Kline Prince William Sound Science Center Zac Hoyt P.O. Box 705 University of Alaska Fairbanks Cordova AK 99574 JC-SFOS 907-424-5800 11120 Glacier Hwy. [email protected] Juneau AK 99801 [email protected] Tim Koeneman Alaska Dept. of Fish & Game Boris Ivanov 917 Sandy Beach Rd. VNIRO Petersburg AK 99833 17, V. Krasnoselskaya 907-772-3422 Moscow 107140 [email protected] RUSSIA [email protected] Kooichi Konishi National Research Institute of Aquaculture Glen Jamieson Nakatsuhamaura 422-1 Dept. of Fisheries & Oceans Nansei Pacific Biological Station Mie 516-0193 Nanaimo BC JAPAN V9R 5K6 CANADA [email protected] 250-256-7223 [email protected] Nikolina Kovatcheva VNIRO Coastal Research Laboratory V. Krasnoselskaya 17 Moscow 107140 RUSSIA [email protected] 850 Participants

Gordon Kruse Jim Meehan University of Alaska Fairbanks NMFS F/ST2 JC-SFOS 1315 East-West Hwy. 11120 Glacier Hwy. Silver Spring MD 20910 Juneau AK 99801-8677 301-713-2363 907-465-8458 [email protected] [email protected] Sue Merkouris Tim Loher Alaska Dept. of Fish & Game University of Washington P.O. Box 25526 School of Fisheries Douglas AK 99802-5526 Box 355020 907-465-6106 Seattle WA 98105 [email protected] 206-547-1035 [email protected] Todd Miller Oregon State University Gustavo Lovrich Dept. of Fisheries Centro Austral de Investigaciones 2030 South Marine Science Drive Cientificas (CADIC) CC-92 Newport OR 97365 Ushuaia 541-867-0100 Tierra del Fuego V9410BFD [email protected] ARGENTINA +54 2901 422310 Adam Moles [email protected] National Marine Fisheries Service Auke Bay Biological Lab Valery Lysenko 11305 Glacier Hwy. TINRO Juneau AK 99801-8626 Naberezhnaya, 18 907-789-6023 Petropavlovsk Kamchatsky [email protected] 683600 RUSSIA [email protected] Holly Moore Alaska Dept. of Fish & Game Kristin Mabry Commercial Fish Alaska Dept. of Fish & Game 333 Raspberry Rd. P.O. Box 25526 Anchorage AK 99518 Juneau AK 99802-5526 907-267-2418 907-465-6111 [email protected] [email protected] Jim Morrison Scott Marshall Dept. of Fisheries & Oceans Alaska Dept. of Fish & Game 3225 Stephenson Point Rd. P.O. Box 240020 Nanaimo BC Douglas AK 99824 V9V 1L3 CANADA 907-465-4260 250-756-7233 [email protected] [email protected]

Takashi Matsuishi Hiroshi Motoh Hokkaido University Kyoto Sea Farming Center Faculty of Fisheries Odasyukuno 3-1-1 Minato-cho Miyazu City Hakodate Kyoto 626-0052 Hokkaido 041-8611 JAPAN JAPAN +81 138 40 8857 Jiro Nagao [email protected] Ehime University Center for Marine Environ. Studies Tony Mecklenburg 3 Bunkyo-cho Point Stephens Research Matsuyama P.O. Box 210307 Ehime 790-8577 Auke Bay AK 99821 JAPAN 907-789-7603 [email protected] [email protected] Crabs in Cold Water Regions: Biology, Management, and Economics 851

Ian Napier Doug Pengilly North Atlantic Fisheries College Alaska Dept. of Fish & Game Port Arthur 211 Mission Rd. Scalloway Kodiak AK 99615 Shetland Isles ZE1 0UN 907-486-2431 UK [email protected] [email protected] Michail Pereladov Einar Nilssen VNIRO University of Tromso V. Krasnoselskaya 17 Norwegian College of Fisheries Moscow 107140 Tromso N-9037 RUSSIA NORWAY [email protected] [email protected] Sara Persselin German Novomodny National Marine Fisheries Service KhoTINRO-Centre 301 Research Court 9 Shevchenko Kodiak AK 99615 Khabarovsk 907-481-1722 RUSSIA [email protected] [email protected] Antan Phillips Shauna Oh Fisheries & Oceans Canada University of California Pacific Biological Station Scripps Inst. of Oceanography 3190 Hammond Bay Rd. 9500 Gilman Drive Nanaimo BC La Jolla CA 92093 V9R 5K6 CANADA 858-822-2708 250-756-7110 [email protected] [email protected]

Bruce Osborne Guy Powell NS Dept. of Agriculture & Fisheries P.O. Box 2285 P.O. Box 2223 Kodiak AK 99615 Halifax, Nova Scotia 907-486-5279 B3T 3C4 CANADA [email protected] 902-424-0352 Martin Robinson Bob Otto Trinity College Dublin National Marine Fisheries Service Dept. of Zoology P.O. Box 1638 Marine Unit Kodiak AK 99615-1638 Dublin 2 907-481-1711 IRELAND [email protected] [email protected]

Wongyu Park Amelie Rondeau University of Alaska Fairbanks Institut Maurice-Lamontagne Fisheries Division Fisheries & Oceans Canada 11120 Glacier Hwy. 850 route de la Mer Juneau AK 99801 CP 1000 Mont-Joli 907-465-6372 Quebec G5H 3Z4 [email protected] CANADA

A.J. Paul Christopher Rooper University of Alaska Fairbanks University of Washington Seward Marine Station School of Fisheries P.O. Box 730 P.O. Box 355020 Seward AK 99664 Seattle WA 98195 206-221-6884 Judy Paul [email protected] University of Alaska Fairbanks Seward Marine Center P.O. Box 730 Seward AK 99664 852 Participants

Gregg Rosenkranz Tom Shirley Alaska Dept. of Fish & Game University of Alaska Fairbanks 211 Mission Rd. Fisheries Division Kodiak AK 99615 11120 Glacier Hwy. [email protected] Juneau AK 99801 907-465-6449 Nicolas Roy [email protected] Maurice Lamontagne Institute 850, Route de la Mer Shareef Siddeek C.P. 1000 Alaska Dept. of Fish & Game Mont-Joli Division of Commercial Fisheries Quebec G5H 3Z4 P.O. Box 25526 CANADA Juneau AK 99802-5526 [email protected] 907-465-6107 [email protected] Janet Rumble Alaska Dept. of Fish & Game Kim Smith P.O. Box 240040 Murdoch University Douglas AK 99824-0020 Div. of Science & Engineering 907-465-4259 South St. [email protected] Murdoch WA 6350 AUSTRALIA Bernard Sainte-Marie [email protected] Maurice Lamontagne Institute Dept. of Fisheries & Oceans Brad Stevens 850, route de la Mer National Marine Fisheries Service CP 1000 Mont-Joli 301 Research Court Quebec G5H 3Z4 Kodiak AK 99615-1638 CANADA 907-481-1726 418-775-0617 [email protected] [email protected] Jan Sundet Herman Savikko Inst. of Fisheries & Aquaculture Alaska Dept. of Fish & Game N-9291 Tromso P.O. Box 25526 NORWAY Juneau AK 99802 [email protected] 907-465-6112 [email protected] Kathy Swiney NMFS Fiona Scurrah 301 Research Court Fisheries & Oceans Canada Kodiak AK 99615 417 2nd Ave. West [email protected] Prince Rupert BC V8J 1G8 CANADA Jim Taggart 250-627-3455 USGS Alaska Biological Science Ctr. [email protected] Glacier Bay Field Station P.O. Box 240009 Lisa Seeb Douglas AK 99824 Alaska Dept. of Fish & Game 907-364-1577 Div of Commercial Fisheries [email protected] 333 Raspberry Rd. Anchorage AK 99518 Shelly Tallack 907-267-2249 P.O. Box 665 [email protected] Stratton Mountain VT 05155 Rich Sewell [email protected] P.O. Box 190914 Anchorage AK 99519 Sherry Tamone 907-562-7837 University of Alaska Southeast 11120 Glacier Hwy. Juneau AK 99801 907-465-6599 [email protected] Crabs in Cold Water Regions: Biology, Management, and Economics 853

Arni Thomson Julio Vinuesa Alaska Crab Coalition CADIC, CONICET 3901 Leary Way NW #6 Av. Malvinas Argentinas s/n Suite 9 Cas. Correo 92 Seattle WA 98107 Ushuaia 9410 206-547-7560 Tierra del Fuego [email protected] ARGENTINA [email protected] Larissa Tolstoganova VNIRO Jonathan Warrenchuk 17 V. Krasnoselskaya St. University of Alaska Fairbanks Moscow 107140 Fisheries Division RUSSIA 11120 Glacier Hwy. [email protected] Juneau AK 99801 [email protected] Charles Trowbridge Alaska Dept. of Fish & Game Leslie Watson 3298 Douglas Place Alaska Dept. of Fish & Game Homer AK 99603 211 Mission Rd. 908-235-8191 Kodiak AK 99615 907-486-1854 Al Tyler [email protected] University of Alaska Fairbanks SFOS Jon Wetzel Fairbanks AK 99775-1090 P.O. Box 959 Homer AK 99603 Anette Ungfors 907-235-7695 Tjarno Marine Biology Lab [email protected] Stromstad 452 96 SWEDEN Dave Witherell +4652668688 North Pacific Fishery Management Council [email protected] 605 W 4th Ave., Suite 306 Anchorage AK 99501 Dan Urban 907-271-2809 Alaska Dept. of Fish & Game [email protected] 211 Mission Rd. Kodiak AK 99615 Astrid Woll 907-486-1849 More Research [email protected] Section of Fisheries P.O. Box 5075 Peter van Tamelen Alesund N-6021 Alaska Dept. of Fish & Game NORWAY Division of Commercial Fisheries [email protected] P.O. Box 25526 Juneau AK 99802-5526 Douglas Woodby 907-465-6129 Alaska Dept. of Fish & Game [email protected] P.O Box 240020 Douglas AK 99824-0020 Ivan Vining 907-465-6115 Alaska Dept. of Fish & Game [email protected] 211 Mission Rd. Kodiak AK 99615 Greg Workman [email protected] Fisheries & Oceans Canada Pacific Biological Station 3190 Hammond Bay Rd. Nanaimo BC V9R 5K6 CANADA [email protected] 854 Participants

Qian-Li Xue The Johns Hopkins University 2024 E. Monument St. Room 2-904 Baltimore MD 21205 410-614-3759 [email protected]

Hiroshi Yamaguchi Hokkaido Central Fisheries Experimental Station Hamanaka 238 Yoichi 046-8555 Hokkaido, JAPAN [email protected]

Stefanie Zaklan University of Alberta Biological Sciences Edmonton, Alberta T6G 2E9 CANADA

Valentina Zheltonozko KamchatNIRO Naberezhnaya 18 Petropavlovsk-Kamchatsky 683600 RUSSIA [email protected]

Jie Zheng Alaska Dept. of Fish & Game P.O. Box 25526 Juneau AK 99802-5526 907-465-6102 [email protected] Crabs in Cold Water Regions: Biology, Management, and Economics 855

Index

A Alitak Bay, Kodiak, Alaska: bitter crab Agnalt, Ann-Lisbeth, 425 syndrome in Chionoecetes bairdi Alaska (Tanner crab) in, 401-403 ADFG Mandatory Shellfish Observer Argentina. See Beagle Channel; San Jorge Program, 693-704 Gulf conclusion, 704 Armstrong, David A., 609 initial program, 694-696 Atlantic Ocean, southwestern. See San North Pacific Fisheries Research Jorge Gulf, Argentina Plan, 698 Auke Bay, Alaska: habitat preferences of overview of program, 702-703 juvenile Chionoecetes bairdi proposal for state-funded program, (Tanner crab) related to oil 698-702 pollution in, 631-642 revisions to program, 696-698 Australia, southern: estimating intermolt mortality of Chionoecetes bairdi duration of Pseudocarcinus gigas (Tanner crab) and Chionoecetes (giant crab) in, 17-28 opilio (snow crab) incidentally caught in Patinopecten B courinus (weathervane Baglin, Raymond E., 305 scallop) fishery in, 717-732 Balzi, Pamela, 283 southeast: restratification of Barents Sea: introduction of Paralithodes Paralithodes camtschaticus camtschaticus (red king crab) to, (red king crab) stock 425-438 assessment areas in,457-473 discussion, 432, 435-436 See also Aleutian Islands; Alitak Bay; materials and methods, 427-430 Auke Bay; Bristol Bay; experiments (tag retention and Frederick Sound; Gulf of juvenile growth), 428, Alaska; Kodiak/Kodiak 429-430 Islands; Prince William Sound; genetic characterization, 428 St. Matthew Island mating behavior, 427-428, 429 Alaskan crabs, checklist of, 5-8 results, 431-432 Aleutian Islands experiment, 429, 431, 433 central: growth of Paralithodes genetics, 431-432, 434 camtschaticus (red king crab) mating, 431 in, 39-50 See also Norway eastern: growth and molting of Beagle Channel, Argentina Lithodes aequispinus (golden growth, maturity, and mating of male king crab) in, 169-187 Lithodes santolla (southern estimating natural mortality of king crab) in, 147-168 Lithodes aequispinus (golden life history of Munida subrugosa king crab) from tag recapture (Galatheid) in, 115-134 data in, 51-75 See also Bering Sea and Aleutian Islands (BSAI) Alinsunurin, Rachel, 537 856 Index

Bering Sea bitter crab disease/syndrome (BCD) and Aleutian Islands (BSAI): estimating in Chionoecetes bairdi (Tanner crab) natural mortality of king crabs (Alitak Bay, Kodiak, Alaska), from tag recapture data, 51-75 401-403 eastern (EBS): in Chionoecetes opilio (snow crab) injuries and aerial exposure to (Newfoundland/Labrador bycatch crabs during continental shelf), 385-400 handling in, 211-212 discussion, 396-398 morphological characteristics of effects on mortality and Chionoecetes hybrids in, recruitment, 397-398 97-113 general distribution, 396-397 observer data in, 537-550 methods, 387-390 historic role of observers, 538 data treatment and analysis, observer database utilization, 389-390 539-548 sampling, 387-389 additional observer data results, 390-396 collections, 544-548 density and abundance area boundaries, 544 relationships, 394-396 gear, 542-543 prevalence and distribution, gear storage, 543-544 389, 390-394 management plans and Blackburn, James E., 213 harvest strategies, Blau, S. Forrest, 39, 51, 169, 213, 225, 305 541 blue king crab. See Paralithodes platypus postseason data analysis, Boutillier, J.A., 439 541-542 Boyle, Larry, 693 preseason fisheries Bristol Bay, Alaska assessment and estimating natural mortality of inseason Paralithodes camtschaticus management, 539- (red king crab) from tag 540 recapture data in, 51-75 stock assessment, 540-541 length-based analysis of Paralithodes program data collection and camtschaticus (red king crab) methods, 538-539 abundance in, 475-494 summary, 548 British Columbia coast, Canada: new spatiotemporal trends in size at fishery for Chionoecetes tanneri maturity of Chionoecetes (grooved Tanner crab) off, 439-456 bairdi (Tanner crab) in, conclusion, 454-455 339-349 methods, 442-446 estimating natural mortality of analysis, 442-446 Lithodes aequispinus (golden sources of mortality, 446 king crab) from tag recapture surveys, 442-444 data in, 51-75 results and discussion, 447-454 mortality of Chionoecetes bairdi biomass estimates, 450, 452 (Tanner crab) as bycatch of distributional trap survey, 444, Patinopecten courinus 448, 450, 451 (weathervane scallop) fishery experimental harvest, 451, 452-453 in, 717-732 phased approach, 454 northwestern: population structure of sources of fishing mortality, 452, Paralithodes platypus (blue 453 king crab) in, 511-520 trawl survey, 447-450 windchill effects on Chionoecetes opilio Bukin, Sergey D., 521 (snow crab) in, 81-96 Burmeister, AnnDorte, 255, 733 Burt, Ryan, 537 Byersdorfer, Susan C., 97, 211, 401 Crabs in Cold Water Regions: Biology, Management, and Economics 857

C Carcinus maenas (European green crab; shore crab) California, northern dispersal on Pacific coast, 561-576 estimating molting probabilities of biological factors, 563-564 female Cancer magister context of regional oceanography, (Dungeness crab) in, 77-80 564-567 sperm plug as indicator of female discussion, 572-573 mating success of Cancer methods, 567-568 magister (Dungeness crab) in, results, 568-571 269-271 in British Columbia, 568-569 Canada. See British Columbia coast oceanographic analyses, 569- Cancer magister (Dungeness crab) 570 estimating molting probabilities of transport mechanisms, 570-571 female (northern California), fishery in Shetland, Scotland, 705, 711 77-80 Cherniawsky, J.Y., 561 habitat use by juvenile, in nursery Chionoecetes bairdi (Tanner crab) estuaries (Pacific coast), bitter crab syndrome in (Alitak Bay, 609-629 Kodiak, Alaska), 401-403 discussion, 621-627 habitat preferences of juvenile, related materials and methods, 610-616 to oil pollution (Auke Bay, data analysis, 616 Alaska), 631-642 habitat characteristics, 614-615 discussion, 640-642 study area, 610-611 methods, 633-635 survey data collection, 611-614 animal collection, 633 results, 616-621 avoidance test protocol, age 0+ results, 618-621 634-635 age 1+ results, 616-618 oiled sediment preparation, 634 comparison of habitat substrate preference, 633 characteristics across results, 635-640 spatial location, 621 response to oil, 635-640 megalopae, relative trophic position of substrate preference, 635 (northern Gulf of Alaska and morphological characteristics of Prince William Sound), hybrids of (eastern Bering 645-649 Sea), 97-113 materials and methods, 645-648 discussion, 110, 112 results and discussion, 648 methods, 99-102 setal stage duration of female adult, results, 102-110 9-15 character scores, 102-103, 104- sperm plug as indicator of female 106 mating success in (northern classification tree, 108, 109-110 California), 269-271 discriminant function analysis, Cancer pagurus (edible crab, European 103, 106-107 edible crab) sources of variation within fishery (Shetland, Scotland), 705, 709- genetic types of 710, 711, 713-714 carapace scores, 108, reproductive capacity 110, 111 morphometrically assessed in mortality of, as bycatch of (Shetland Island), 405-423 Patinopecten courinus discussion, 413-417, 419-421 (weathervane scallop) fishery materials and methods, 408-411 (Bering Sea), 717-732 results, 412 data and methods, 718-719 abdomen width, 412, 415, 416, discussion, 730-731 417, 418-419 results, 719-730 sexual dimorphism and chela generalized linear model allometry, 412, 413, (GLM), 726-730 414 858 Index

Chionoecetes bairdi, mortality of Chionoecetes opilio, female reproductive (continued) condition (continued) graphical analysis and results, 260-263 summary statistics, annual changes in color of 719-726 brood, 260 spatiotemporal trends in size at clutch and ovary weight in maturity of (eastern Bering subsamples, 261, 262, Sea), 339-349 263 methods, 340-341 number of eggs per clutch, results and discussion, 341-348 261-262, 264, 265 survival of, tagged with Floy tags, weather and temperature 551-560 conditions, 260, 261 discussion, 556-558 male distribution and demography materials and methods, 552-554 (Newfoundland/Labrador results, 554-556 continental shelf), 577-594 Chionoecetes japonicus (red snow crab): discussion, 587-592 larval development of (Osaka, effects of depth on Japan), 135-146 temperature, 589-592 discussion, 143-145 trends in distribution and size materials and methods, 136 composition, 587-589 results, 136-143 methods, 578-581 crab 1, 142-143, 144 data collection, 578-580 megalopa, 139-142, 143 data treatment and analysis, zoea 1, 137-138 581 zoea 2, 138-139 results, 581-587 Chionoecetes opilio (snow crab) distribution, 581-583 bitter crab disease in (Newfoundland/ size composition, 583-586 Labrador continental shelf), size segregation by depth and 385-400 temperature, 586-587 discussion, 396-398 morphological characteristics of effects on mortality and hybrids of (eastern Bering recruitment, 397-398 Sea), 97-113 general distribution, 396-397 discussion, 110, 112 methods, 387-390 methods, 99-102 data treatment and analysis, results, 102-110 389-390 character scores, 102-103, 104- sampling, 387-389 106 results, 390-396 classification tree, 108, 109-110 density and abundance discriminant function analysis, relationships, 394-396 103, 106-107 prevalence and distribution, sources of variation within 389, 390-394 genetic types of female reproductive condition (west carapace scores, 108, Greenland), 255-267 110, 111 conclusion, 265-266 mortality of, as bycatch of discussion, 263-265 Patinopecten courinus egg stages, 263-264 (weathervane scallop) fishery ovaries and clutch, 264-265 (Bering Sea), 717-732 materials and methods, 257-260 data and methods, 718-719 data analysis, 260 discussion, 730-731 study area and sampling results, 719-730 procedure, 257-260 generalized linear model subsampling of females and (GLM), 726-730 laboratory processing, graphical analysis and 260 summary statistics, 719-726 Crabs in Cold Water Regions: Biology, Management, and Economics 859

Chionoecetes opilio (snow crab) (continued) E spatiotemporal trends in (eastern edible crab. See Cancer pagurus Bering Sea), 339-349 Erimacrus isenbeckii (horsehair crab; methods, 340-341 Japanese hair crab) results and discussion, 341-348 population assessment using length- windchill effects on, 81-96 based analysis for (eastern discussion, 90-94 Hokkaido, Japan), 495-509 materials and methods, 84-86 discussion, 504-508 statistical analysis, 86 methods, 496-504 windchill treatments, 85-86 data, 496-498 results, 86-90 LPA model, 498-502 autonomy, 87-88, 89 parameter estimation, 502-504 mortality, 86-87, 88 results, 504 righting response, 88, 90, 91 trap/trawl data for (Tatar Strait, Chionoecetes tanneri (grooved Tanner southern, Russia), 522-525, crab): new fishery for (British 526-527, 532-535 Columbia coast, Canada), 439-456 European edible crab. See Cancer pagurus conclusion, 454-455 European green crab. See Carcinus maenas methods, 442-446 analysis, 442-446 F sources of mortality, 446 Farestveit, Eva, 425 surveys, 442-444 Foreman, M.G.G., 561 results and discussion, 447-454 Frederick Sound, Alaska: movement and biomass estimates, 450, 452 habitat utilization by Lithodes distributional trap survey, 444, aequispinus (golden king crab) in, 448, 450, 451 595-608 experimental harvest, 451, 452-453 phased approach, 454 G sources of fishing mortality, 452, 453 Gardner, Caleb, 17 trawl survey, 447-450 giant crab. See Pseudocarcinus gigas Chizzini, Alejandro, 115 golden king crab. See Lithodes aequispinus Clark, John E., 457 Grays Harbor, Washington. See Washington Colbourne, Eugene B., 577 coast Coos Bay, Oregon. See Oregon coast green crab, European. See Carcinus maenus crustaceans: estimating duration of molt Greenland stages in, 351-365 southeast: Lithodes maja (northern discussion, 361, 363-365 stone crab) at, 733-749 estimator development, 352-358 west (Disko Bay and Sisimiut): Buchholz (1991) method, 353-355 reproductive condition of proposed new methods, 356-358 mature female Chionoecetes results, 360-361 opilio (snow crab) in, 255-267 days to molt experiments, 361, 362 Gulf of Alaska: relative trophic position of days to stage experiments, 360-361 Cancer magister (Dungeness simulation methods, 358-360 crab) megalopae, in, 645-649 days to molt experiments, 359 Gunderson, Donald R., 609 days to stage experiments, 358-359 H D hair crab, Japanese. See Erimacrus Dawe, Earl G., 385, 577 isenbeckii Deadman Reach. See Alaska, southeast Hanasaki king crab. See Paralithodes Disko Bay, west Greenland. See Greenland, brevipes west Hankin, David G., 9, 77, 269, 351 Dungeness crab. See Cancer magister Hapalogastrinae subfamily. See Lithodidae family 860 Index

Heijnis, Hendrik, 17 Kodiak/Kodiak Island, Alaska helmet crab. See Telmessus cheiragonus Alitak Bay: bitter crab syndrome in Hematodinium species. See bitter crab Chionoecetes bairdi (Tanner disease crab) in, 401-403 Hinkley, Sandy, 457 archipelago: mating pairs of Hjelset, Ann Merete, 681 Paralithodes camtschaticus Hokkaido, Japan (red king crab) in, 225-245 eastern: population assessment for fecundity and clutch fullness of Erimacrus isenbeckii Paralithodes camtschaticus (Japanese hair crab) using (red king crab) at, 305-321 length-based analysis, 495-509 Management Area: female size at southern: reproductive cycle of maturity of Paralithodes Telmessus cheiragonus camtschaticus (red king crab) (helmet crab) in, 323-337 in, 213-224 horsehair crab. See Erimacrus isenbeckii discussion, 220-222 Hoyt, Zachary N., 595 methods, 214-218 results, 218-220 I Koeneman, Timothy, 457 Ishikawa, Manabu, 189 Konishi, Kooichi, 135 Ivanov, Boris G., 651 Kovatcheva, Nikolina, 273 Kruse, Gordon H., 367, 475 J Kuril Islands, Russia: trap catch data for Jadamec, Luke, 97 Lithodes aequispinus (golden king Jamieson, G.S., 561 crab) in, 525-535 Japan. See Hokkaido; Osaka Japanese hair crab. See Erimacrus L isenbeckii Labrador/Newfoundland continental shelf Jenkinson, Andrew, 17 bitter crab disease in Chionoecetes Johnson, B. Alan, 305 opilio (snow crab) at, 385-400 Jørstad, Knut E., 425 distribution and demography of Chionoecetes opilio (snow K crab) males at, 577-594 Kamchatka coast/shelf, western, Russia Levings, C.D., 561 stock management problems and Lithodes aequispinus (golden king crab) research of Paralithodes Benedict, 1895: correct spelling and camtschaticus (red king crab) publication data of, 1-3 in, 651-680 estimating natural mortality of, from trap/trawl catch data for Paralithodes tag recapture data (Aleutian camtschaticus (red king crab) Islands, Alaska), 51-75 on, 522-525, 526-527, 532-535 discussion, 69-72 Kanno, Yasuji, 495 materials and methods, 53-61 king crab. See Lithodes species; development of M estimator, Paralithodes species 55-61 king crab, blue. See Paralithodes platypus tag releases, 53-55 king crab, golden. See Lithodes aequispinus results, 61-69 king crab, Hanasaki. See Paralithodes growth and molting of (eastern Aleutian brevipes Islands, Alaska), 169-187 king crab, red. See Paralithodes discussion, 182-185 camtschaticus methods, 170-172 king crab, southern. See Lithodes santolla results, 173-182 Kittaka, Jiro, 189 female growth, molting Kline, Thomas C., 645 probability, and reproductive cycle, 178-182 Crabs in Cold Water Regions: Biology, Management, and Economics 861

Lithodes aequispinus, growth and molting Lithodes santolla (southern king crab) (continued) (continued) male growth and molting morphometric and behavioral probability, 173-178 maturity, 155, 157- movement and habitat utilization by 158, 160, 161 (Frederick Sound, Alaska), reproductive biology of (San Jorge 595-608 Gulf, Argentina), 283-304 discussion, 604-606 discussion, 298-301 movements, 605-606 materials and methods, 284-287 observations, 604-605 results, 287-298 materials and methods, 597-599 embryogenesis and egg size, results, 600-604 290-291 movements, 601-604 fecundity, 292, 293 observations, 600-601 female molting, 288, 289 trap catch data for (Kuril Islands, reproductive cycle, 287-288 Russia), 525-535 reproductive structure, 288- Lithodes maja (northern stone crab): 290, 291 occurrence of (southeast sex proportion, relative Greenland), 733-749 abundance, and conclusions, 747 movement, 292, 294- discussion, 745-747 298 catch rates, 745-746 Lithodidae family (Pacific Ocean, north), processing and meat yield, 747 751-845 reproduction and fecundity, 746-747 conclusion, 761-762 materials and methods, 735-737 evolutionary relationships, 752, 763 biological data, 736 fisheries, 821-823 catch data, 735-736, 739, 741 life history traits, 799-814 gear, 735 parasites and commensals, 754-755, processing and market testing, 736 818-820 study area, 735, 736, 737 predator/prey relationships, 815-817 results, 737-745 subfamilies, 755-761 catch rates, 737-740 Hapalogastrinae subfamily, 755-756 processing and market testing, life history traits, 812-814 741-744 predator/prey relationships, reproduction and fecundity, 740- 817 741, 743-745 parasites and commensals, 820 sex and size distribution, 740, 741, summary data, 796-798 742 Lithodinae subfamily, 756-761 Lithodes santolla (southern king crab) adaptations, 759 growth, maturity, and mating of male behaviors, 758 (Beagle Channel, Argentina), fisheries, 759-761 147-168 life cycle, 756-758 discussion, 160, 162-164 life history traits, 799-812 materials and methods, 151-154 parasites and commensals, growth, 152-153 818-820 morphometric maturity, predator/prey relationships, 153-154 815-817 study site and sampling, summary data, 779-796 151-152 results, 154-160 sperm morphology, 754 growth increments, 154-155, summary data, 764, 779-798 156, 157 taxonomic keys, 777-778 molting frequency, 155, 158, Lithodinae subfamily. See Lithodidae family 159 Lovrich, Gustavo A., 115, 147 862 Index

M Norway fishery management and bycatch of Matsuishi, Takashi, 495 Paralithodes camtschaticus Matsumoto, Toshie, 135 (red king crab) in, 681-692 Miljutin, D.M., 511 See also Barents Sea Miller, Todd W., 9 Moles, Adam, 631 O molt stages in crustaceans, estimating, 351-365 O’Clair, Charles E., 595 discussion, 361, 363-365 Oh, Shauna J., 269 estimator development, 352-358 Okhotsk Sea. See Kamchatka coast/shelf, Buchholz (1991) method, 353-355 western, Russia proposed new methods, 356-358 Olsen, Steinar, 425 results, 360-361 Oregon coast: habitat use by juvenile days to molt experiments, 361, 362 Cancer magister (Dungeness days to stage experiments, 360-361 crab) in nursery estuaries of, simulation methods, 358-360 609-629 days to molt experiments, 359 Osaka, Japan: larval development of days to stage experiments, 358-359 Chionoecetes japonicus (red snow Moore, Holly, 51, 537 crab), 135-146 Moscow, Russia: rearing zoeae and Otto, Robert S., 339 glaucothoe of Paralithodes camtschaticus (red king crab) in P recycling water (CRAS) system in, Pacific coast 273-282 dispersal of Carcinus maenas Munehara, Hiroyuki, 323 (European green crab) on, Munida gregaria. See Munida subrugosa 561-576 Munida subrugosa: life history of (Beagle habitat use by juvenile Cancer Channel, Argentina), 115-134 magister (Dungeness crab) in discussion, 128-131 nursery estuaries of, 609-629 materials and methods, 118-121 See also British Columbia coast, density and biomass, 119 Canada; California, northern; maturity size, 119 Oregon coast; Washington natural diet, 120-121 coast reproductive cycle, 119-120 Pacific Ocean, north: Lithodidae family in, study site and sampling, 118-119 751-845 results, 121-128 Paralithodes brevipes (Hanasaki king crab): density and biomass, 121, 122 larval culture of, 189-209 fecundity, 124 discussion, 204-206 feeding habits, 124, 127-128 materials and methods, 190-194 gonadal and morphometric experiment with enriched Artemia maturity, 121, 123 nauplii, 192-193 reproductive cycle, 121, 124, 125, fat analysis of cultured and 126 enriched Thalassiosira and hatched Artemia, N 193-194 Nagao, Jiro, 323 ordinal foods, with 190-192 Napier, Ian R., 705 statistical procedures, 193 Necora puber (velvet crab): fishery results, 194-204 (Shetland, Scotland), 705, 711, larval and postlarval culture, 194, 713-714 196-197 Newfoundland/Labrador continental shelf lipid content and fatty acid bitter crab disease in Chionoecetes composition of cultured opilio (snow crab) at, 385-400 Thalassiosira, 195, 199- Nizyaev, Sergey A., 521 202 survival rate, 195, 203 Crabs in Cold Water Regions: Biology, Management, and Economics 863

Paralithodes brevipes, larval culture Paralithodes camtschaticus, fishery (continued) management (continued) Thalassiosira culture, 194-195, 198 gear and fishing regulations, zoea and glaucothoe stages, 195, 683-685 202, 204 participation, 683 Paralithodes camtschaticus (red king crab) growth of (central Aleutian Islands), acoustical behavior in, 247-254 39-50 discussion, 252, 254 discussion, 46, 48-49 materials and methods, 248-249 methods, 40-42 fishing experiments, 249 results, 42-46, 47, 48 reaction studies, 248-249 habitat preferences of juvenile, related sound studies, 248 to oil pollution (Auke Bay, results, 250-252 Alaska), 631-642 behavior, 250 discussion, 640-642 fishing experiments, 252 methods, 633-635 reactions to natural crab sound animal collection, 633 emission, 252, 253 avoidance test protocol, sounds, 250, 251 634-635 estimating natural mortality of, from oiled sediment preparation, 634 tag recapture data (Bristol substrate preference, 633 Bay, Alaska), 51-75 results, 635-640 discussion, 69-72 response to oil, 635-640 materials and methods, 53-61 substrate preference, 635 development of M estimator, introduction of (Barents Sea), 425-438 55-61 discussion, 432, 435-436 tag releases, 53-55 materials and methods, 427-430 results, 61-69 experiments (tag retention and fecundity and clutch fullness of juvenile growth), 428, (Kodiak Island, Alaska), 429-430 305-321 genetic characterization, 428 discussion, 318-319 mating behavior, 427-428, 429 materials and methods, 306, 308- results, 431-432 310 experiment, 431 clutch fullness agreement, genetics, 431-432 309-310 mating, 431 embryo numbers, 308-309 larval culture of, 189-209 sample collection, 306, 308 discussion, 204-206 results, 310-317 materials and methods, 190-194 fecundity estimation, 307, 310- experiment with enriched 312, 313 Artemia nauplii, pairwise agreement among 192-193 raters, 312, 314-317 fat analysis of cultured and female size at maturity of (Kodiak enriched Thalassiosira Management Area, Alaska), and hatched Artemia, 213-224 193-194 discussion, 220-222 ordinal foods, with 190-192 methods, 214-218 statistical procedures, 193 results, 218-220 results, 194-204 fishery management and bycatch of larval and postlarval culture, (Norway), 681-692 194, 196-197 bycatch, 689 lipid content and fatty acid discussion, 690-691 composition of fishery, 683-689 cultured Thalassiosira, catch quotas and CPUE, 685-689 195, 199-202 fishing areas, 685 survival rate, 195 864 Index

Paralithodes camtschaticus, larval culture Paralithodes camtschaticus (red king crab) (continued) (continued) Thalassiosira culture, 194-195, restratification of stock assessment 198 areas of (southeast Alaska), zoea and glaucothoe stages, 457-473 195, 202, 204 discussion, 470-471 length-based analysis of abundance of methods, 458-465 (Bristol Bay, Alaska), 475-494 Deadman Reach example, 465- methods, 476-481 470 data, 479 restratification, 464-465 female population model, 479 spatial analysis, 461, 463-464 male population model, survey methods, 458-461, 462 476-479 stock management problems and parameter estimation, 480 research of (western S-R models, 481 Kamchatka shelf, Russia), results and discussion, 481-492 651-680 management implications, discussion and conclusions, 673- 488-492 677 population abundance, 481-486 fishery biology, 659-667 S-R relationship, 486-488 problems, 666-667 mating pairs of (Kodiak archipelago, research concepts, 661-666 Alaska), 225-245 research history, 660-661 discussion, 238-242 fishery management, 668-673 methods, 227-231 federal level, 669-671 results, 232-238 history, 668 habitat and depth of collected international level, 668-669 grasping pairs, 228- regional level, 671-673 230, 232 fisheries, 652-659 monthly changes in size of dynamics, 657-659 mating females and harvesting areas, 656-657 males, 236-238 history, 652-656 observations by scuba divers, trap/trawl catch data for (western 232-233 Kamchatka coast, Russia), seasonal presence of grasping 522-525, 526-527, 532-535 pairs and seasonal Paralithodes platypus (blue king crab) changes in diver CPUE, abundance assessment of, using catch- 228-229, 233, 234 survey model (St. Matthew size and shell-age composition Island, Alaska), 367-384 of females and males, abundance projections, 378-379 233, 235 change in natural mortality, size relationships of males and 370-372 females within pairs, discussion, 379-382 235-236 model assessment, 372-377 molting of, observed by time-lapse data, 373 video, 29-37 four-stage model, 372-373 discussion, 35-36 model fit, 374-377 materials and methods, 30-31, 32 parameter estimation, 373-374 results, 31, 33-35 estimating natural mortality of, from rearing zoeae and glaucothoe of, in tag recapture data (St. recycling water (CRAS) system Matthew Island, Alaska), (Moscow, Russia), 273-282 51-75 discussion, 279-281 discussion, 69-72 materials and methods, 274-275, materials and methods, 53-61 276 development of M estimator, results, 275-279 55-61 Crabs in Cold Water Regions: Biology, Management, and Economics 865

Paralithodes platypus, estimating natural Rudra, Hari, 425 mortality (continued) Russia. See Barents Sea; Kamchatka coast; tag releases, 53-55 Kuril Islands; Moscow; Sakhalin results, 61-69 coast; Tatar Strait, southern population structure of (northwestern Bering Sea), 511-520 S discussion, 518-520 St. Matthew Island, Alaska materials and methods, 512-514 abundance assessment of Paralithodes results, 514-518 platypus (blue king crab) trap catch data for (Sakhalin coast, using catch-survey model, Russia), 526, 532-535 367-384 Patinopecten courinus (weathervane estimating natural mortality of scallop): mortality of Paralithodes platypus (blue Chionoecetes bairdi (Tanner crab) king crab) from tag recapture as bycatch of (Bering Sea), data, 51-75 717-732 Sakhalin coast, Russia: trap catch data for data and methods, 718-719 Paralithodes platypus (blue king discussion, 730-731 crab) on, 526, 532-535 results, 719-730 San Jorge Gulf, Argentina: reproductive generalized linear model (GLM), biology of Lithodes santolla 726-730 (southern king crab) in, 283-304 graphical analysis and summary Schwenzfeier, Mary, 537, 693 statistics, 719-726 Scotland. See Shetland/Shetland Island Pengilly, Douglas, 39, 97, 169, 213, 225, 339 Scurrah, F.E., 439 Pereladov, M.V., 511 Sea of Japan. See Tatar Strait, southern, Phillips, A.C., 439 Russia Powell, Guy C., 225 shellfish: ADFG Mandatory Observer Prince William Sound, Alaska: trophic Program of (Alaska), 693-704 position of Cancer magister conclusion, 704 (Dungeness crab) megalopae in, initial program, 694-696 645-649 North Pacific Fisheries Research Plan, Pseudocarcinus gigas (giant crab): 698 estimating intermolt duration of overview of program, 702-703 (southern Australia), 17-28 proposal for state-funded program, discussion, 24, 26 698-702 methods, 20-22 revisions to program, 696-698 proportion of females reproducing, Shetland/Shetland Island, Scotland 20-21 fisheries in, 705-716 test of assumptions of radiometric current, 711-714 aging, 21-22 development, 709-711 results, 22-24 history, 706-709 proportion of females reproducing, management, 714-716 22, 23 reproductive capacity test of assumption of radiometric morphometrically assessed in aging, 22, 24, 25, 26 Cancer pagurus in, 405-423 Shirley, Thomas C., 1, 81, 595 R shore crab. See Carcinus maenus red king crab. See Paralithodes Siddeek, M.S.M., 51 camtschaticus Sisimiut, west Greenland. See Greenland, red snow crab. See Chionoecetes japonicus; west C. tanneri Smith, Barry D., 147 Romero, M. Carolina, 115 snow crab. See Chionoecetes opilio Rooper, Christopher N., 609 Stevens, Bradley G., 5, 29, 189, 551 Rosenkranz, Gregg E., 717 866 Index

stock assessment using catch-survey trap catch data, assessing crab resources model, 367-384 based on (continued) abundance projections, 378-379 effect of soak time on trap catches, change in natural mortality, 368, 370-372 526 discussion, 379-382 results, 526-532 model assessment, 372-377 comparing trap types, 527 data, 373 comparing trawl and trap data, four-stage model, 372-373 526-527 model fit, 374-377 effect of soak time on trap catches, parameter estimation, 373-374 527-532 stone crab, northern. See Lithodes maja Tsujimoto, Ryo, 135 Stone, Robert P., 595, 631 Sundet, Jan H., 681 U Ueda, Yuji, 495 T Urban, Daniel, 97, 401 Tallack, Shelly M.L., 405 Usujiri, Japan. See Hokkaido, Japan, Tanner crab. See Chionoecetes bairdi southern Tanner crab, grooved. See Chionoecetes tanneri V Tapella, Federico, 115 Varangerfjord, Norway. See Barents Sea Tatar Strait, southern, Russia: trap/trawl velvet crab. See Necora puber data for Erimacrus isenbeckii Vining, Ivan, 39 (horsehair crab) in, 522-525, 526- Vinuesa, Julio H., 147, 283 527, 532-535 Telmessus cheiragonus (helmet crab): W reproductive cycle of, 323-337 Warrenchuk, Jonathan J., 81, 595 discussion, 334-336 Washington coast: habitat use by juvenile materials and methods, 324-325, 326 Cancer magister (Dungeness results, 325, 327-337 crab) in nursery estuaries of, gonadal maturation, 327, 328-331 609-629 gonad anatomy and histology, 325 Watson, Leslie J., 51, 169 minimum mature size, 327, 332 West Coast. See Pacific coast molting frequency in females, 327, Willapa Bay, Washington. See Washington 334 coast monthly change of gonadal Woll, Astrid K., 733 maturity and GSI value, Workman, G.D., 439 327, 333 Teshima, Shin-ichi, 189 X Tolstoganova, Larissa K., 247 Xue, Qian-Li, 77 Tracy, Donn A., 211 trap catch data, assessing crab resources Y based on, 521-536 discussion, 532-535 Yamaguchi, Hiroshi, 495 materials and methods, 522-526 Yaquina Bay, Oregon. See Oregon coast comparing trap catches with environmental variation Z (Lithodes aequispinus, Zaklan, S.D., 751 Paralithodes platypus), Zheng, Jie, 367, 475 526 comparing trawl and trap data, 522-525 comparing two trap type catches (Lithodes aequispinus), 522-525