BIOLOGY, MANAGEMENT, AND ECONOMICS

A.J. Paul, Earl G. Dawe, Robert Elner, Glen S. Jamieson, Gordon H. Kruse, Robert S. Otto, Bernard Sainte-Marie, Thomas C. Shirley, and Douglas Woodby, Editors

Proceedings of the symposium Crab2001, in Cold Water Regions: Biology, Management, and Economics January 17-20, 2001, Anchorage, , USA

University of Alaska Sea Grant College Program AK-SG-02-01

Price: $40.00 Elmer E. Rasmuson Library Cataloging-In-Publication Data

Crabs in cold water regions : biology, management, and economics / Editors: A.J. Paul … [et al.]. Fairbanks, Alaska : University of Alaska Sea Grant, [2002].

876 p. ; cm. – (Lowell Wakefield Fisheries Symposium ; [19th]), (University of Alaska Sea Grant College Program ; AK-SG-02-01)

Note: “... Proceedings of the symposium Crab2001, Crabs in cold water regions: biology, management, and economics, January 17-20, 2001, Anchorage, Alaska, USA.”

Includes bibliographical references and index

1. Crabs—Congresses. 2. fisheries—Congresses. I. Title. II. Paul, A. J. III. Series: Lowell Wakefield Fisheries Symposium series ; 19th. IV. Series: Alaska Sea Grant College Program report ; AK-SG-02-01.

QL444.M33 C73 2002

ISBN: 1-56612-077-2

Citation for this volume is: 2002. A.J. Paul, E.G. Dawe, R. Elner, G.S. Jamieson, G.H. Kruse, R.S. Otto, B. Sainte-Marie, T.C. Shirley, and D. Woodby (eds.). Crabs in Cold Water Regions: Biology, Management, and Economics. University of Alaska Sea Grant, AK-SG- 02-01, Fairbanks. 876 pp.

Credits This book is published by the University of Alaska Sea Grant College Program, which is cooperatively supported by the U.S. Department of Commerce, NOAA National Sea Grant Office, grant no. NA86RG-0050, project A/161-01; and by University of Alaska Fairbanks with state funds. University of Alaska is an affirmative action/equal opportunity institu- tion. Sea Grant is a unique partnership with public and private sectors combining re- search, education, and technology transfer for public service. This national network of universities meets changing environmental and economic needs of people in our coastal, ocean, and Great Lakes regions.

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U E (907) 474-6707 Fax (907) 474-6285 S C D R E E P M A M RT O MENT OF C http://www.uaf.edu/seagrant/ About the Symposium Crab is one of the world’s most valuable marine consumables, especially to Alaska. So it is not surprising that the topic of crab has been addressed more often than any other by the Lowell Wakefield Symposium series, each time at the request of resource managers and researchers. Crab2001, Crabs in Cold Water Regions: Biology, Management, and Economics, held January 17-20, 2001 in Anchorage, Alaska, was the sixth crab symposium in the series (1982, 1984, 1985, 1989, 1995, and 2001). The year for the Crab2001 symposium had been “selected” six years earlier by participants at the 1995 Wakefield symposium on high latitude crabs. The symposium was organized and coordinated by Brenda Baxter, University of Alaska Sea Grant Program, with the assistance of the orga- nizing committee. Committee members are: Earl Dawe, Department of Fisheries and Oceans, Canada; Glen Jamieson, Department of Fisheries and Oceans, Canada; Gordon Kruse, University of Alaska Fairbanks, School of Fisheries and Ocean Sciences (formerly of Alaska Department of Fish and Game); Bob Otto, U.S. National Marine Fisheries Service, Alaska Fisher- ies Science Center; A.J. Paul, University of Alaska Fairbanks, Institute of Marine Science; and Dave Witherell, North Pacific Fishery Management Council. Symposium sponsors are: University of Alaska Sea Grant College Pro- gram; Alaska Department of Fish and Game; North Pacific Fishery Manage- ment Council; U.S. National Marine Fisheries Service; and Wakefield Endowment, University of Alaska Foundation.

The Lowell Wakefield Symposium Series The University of Alaska Sea Grant College Program has been sponsoring and coordinating the Lowell Wakefield Fisheries Symposium series since 1982. These meetings are a forum for information exchange in biology, management, economics, and processing of various fish species and com- plexes as well as an opportunity for scientists from high latitude coun- tries to meet informally and discuss their work. Lowell Wakefield was the founder of the Alaska industry. He recognized two major ingredients necessary for the king crab fishery to survive—ensuring that a quality product be made available to the con- sumer, and that a viable fishery can be maintained only through sound management practices based on the best scientific data available. Lowell Wakefield and Wakefield played important roles in the develop- ment and implementation of quality control legislation, in the prepara- tion of fishing regulations for Alaska waters, and in drafting international

ix agreements for the high seas. Toward the end of his life, Lowell Wakefield joined the faculty of the University of Alaska as an adjunct professor of fisheries where he influenced the early directions of the university’s Sea Grant Program. This symposium series is named in honor of Lowell Wake- field and his many contributions to Alaska’s fisheries. Three Wakefield symposia are planned for 2003-2005.

Proceedings Acknowledgments This publication presents 53 symposium papers. Each full-length paper was reviewed by two peer reviewers, extended abstracts had one review each, and papers were revised according to recommendations by associ- ate editors who generously donated their time and expertise: A.J. Paul, Earl G. Dawe, Robert Elner, Glen S. Jamieson, Gordon H. Kruse, Robert S. Otto, Bernard Sainte-Marie, Thomas C. Shirley, and Douglas Woodby. The first two papers, by T.C. Shirley and B.G. Stevens, and the last, by S.D. Zaklan, were not presented at the symposium; the editors chose to include them in the book. Thanks go to the authors of all 53 contribu- tions. Many thanks to the following people who reviewed one or more manu- scripts for this book: Klaus Anger, Dave Armstrong, Celine Audet, David Barnard, Jim Blackburn, Jim Boutillier, Forrest Bowers, Ryan Burt, Larry Byrne, Alan Campbell, John Clark, J. Crain, Paula Cullenberg, Braxton Dew, Bill Donaldson, Rejean Dufour, Bob Elner, Darryl Felder, Richard Forward, Caleb Gardner, Skip Gish, Don Gunderson, David Hankin, Gretchen Har- rington, Marcel Hebert, S.Y. Hong, Luke Jadamec, Glen Jamieson, Stephen Jewett, B. Alan Johnson, Knut Jørstad, Jiro Kittaka, Tom Kline, Kooichi Konishi, Gordon Kruse, Andrew Levings, Gustavo Lovrich, Patsy A. McLaugh- lin, Tony Mecklenburg, G.A. Messick, Bob Miller, Adam Moles, Frank Morado, Mikio Moriyasu, Holly Moore, Hiroyuki Munehara, Jiro Nagao, Peter Ng, Chuck O’Clair, Steinar Olsen, Wongyu Park, A.J. Paul, Judy Paul, Doug Pengilly, Ian Potter, Martin Robinson, Amelie Rondeau, Christopher Rooper, Gregg Rosenkranz, Janet Rumble, Mary Schwenzfeier, Tom Shirley, Shareef Siddeek, Barry Smith, Brad Stevens, Jan Sundet, Kathy Swiney, Arnie Thomson, Shelly Tallack, Dave Taylor, Donn Tracy, Oliver Tully, Al Tyler, Sherry Tamone, Federico Tapella, John Tremblay, Dan Urban, Peter van Tamelen, Julio Vinuesa, Ivan Vining, Elmer Wade, Jonathon Warrenchuk, Leslie Watson, Greg Workman, Hiroshi Yamaguchi, Zane Zhang, and Jie Zheng. Copy editing is by Kitty Mecklenburg of Pt. Stephens Research Associ- ates, Auke Bay, Alaska; and Sue Keller, University of Alaska Sea Grant. Layout and format are by Kathy Kurtenbach, and cover design is by Tatiana Piatanova, both of University of Alaska Sea Grant.

x Crabs in Cold Water Regions: Biology, Management, and Economics 855

Index

A Alitak Bay, Kodiak, Alaska: bitter crab Agnalt, Ann-Lisbeth, 425 syndrome in 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 () 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 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 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, () 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 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, (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 : 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