Dungeness crab {Cancer magister) recruitment variability and Ekman transport of larvae

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1 ICES mar. Sei. Symp., 199: Dungeness crab {Cancer magister) recruitment variability and Ekman transport of larvae Robert A. McConnaughey, David A. Armstrong, and Barbara M. Hickey McConnaughey, R. A., Armstrong, D. A., and Hickey, B. M Dungenesscrab (Cancer magister) recruitment variability and Ekman transport of larvae. - ICES mar. Sei. Symp., 199: Commercial stocks of the Dungeness crab (Cancer magister) fluctuate widely along the Pacific coast of the United States. Previous attempts to explain this variability have focused on the pelagic larval phase and the environmental factors that may affect survival and dispersal of this stage. However, these efforts have been largely unsuccessful because, at least in part, commercial landings data are used to measure year-class strength and correspondingly long time-lags (3-4 years) are required. In addition, studies concerned with the advective transport of larvae have not corrected calculated winds for wind veering near the coast (we have). In this study, we compare the abundance of newly settled crab from trawl surveys along the southern Washington coast ( ) with wind-driven (Ekman) transport during the preceding larval periods. As such, the operative time-lag is on the order of months. Results suggest that strong landward transport is conducive to heavy C. magister settlement. This condition will tend to minimize cross-shelf advective loss of larvae while promoting retention near suitable juvenile and adult habitat. Strong alongshore Ekman transport, on the other hand, was apparently detrimental to C. magister recruitment. This latter effect has not been described previously and represents a significant departure from conventional thinking about the dynamics of C. magister larvae (which focuses almost exclusively on the cross-shelf component of the coastal circulation). Moreover, our analyses do not support the paradigm that C. magister larvae are transported progressively seaward through ontogeny, since there is no evidence of sustained westward Ekman transport during the six larval periods investigated. Robert A. McConnaughey: School o f Fisheries, University o f Washington, Seattle, W A 98195, USA. Current address: A laska Fisheries Science Center, National Marine Fisheries Service, 7600 Sand Point Way N. E., Seattle, W A , USA. David A. Arm strong: School o f Fisheries, University o f Washington, Seattle, W A 98195, USA. Barbara M. Hickey: School o f Oceanography, University o f Washington, Seattle, W A 98195, USA. [tel: (+1) , fax: (+1) ], Introduction The Dungeness crab (Cancer magister) is widely distributed in the eastern Pacific Ocean and supports important commercial fisheries characterized by highly variable and cyclical landings (Methot, 1986; Botsford et al., 1989). Its complex life history is typical of marine invertebrates with meroplanktonic larvae and a broadcast reproductive strategy. In coastal Washington waters, mating occurs primarily in May-June, eggs are extruded in September-October, larvae are released during January to March (Cleaver, 1949). Larval ontogeny requires days and consists of five zoeal stages and the megalopa stage (Poole, 1966; Reed, 1969; Lough, 1976). Settlement to the benthos occurs primarily during May-June followed, perhaps, by a second smaller pulse later that same year (Stevens and Armstrong, 1984; Gunderson et al., 1990; unpublished data). The rate of growth is highly variable (Botsford, 1984; Orensanz and Gallucci, 1988) and it is generally held that entry into the fishery occurs three to four years after hatching. Previous attempts to explain substantial interannual variability in C. magister abundance have been largely unsuccessful (reviewed by Hankin, 1985 ; Botsford et al., 1989). Most have investigated time-lagged relationships between physical features of the environment during the planktonic larval phase and commercial fishery landings (Peterson, 1973; Botsford and Wickham, 1975; Wild, 1980, 1983; Love and Westphal, 1981; Johnson et al., 1986). Other studies considered the effects of biotic

2 168 R. A. McConnaughey, D. A. Armstrong, and B. M. Hickey ICES mar. Sei. Symp., 199 (1995) 47 00'N Willapa Bay depth isobath (m) 46 30'N 'W "W Figure 1. Study area along the southern Washington coast showing subsystem boundaries, nearshore transect lines, trawl stations (filled circles) and the site (+ ) selected for Ekman transport data used in the analyses. Dashed lines indicate stratum boundaries which, for the nearshore, are positioned along the mean lower low water line and the 14m (7.5fm), 41 m (22.5fm), and 59m (32.5 fm) contours. factors and were also inconclusive (Botsford and Wickham, 1978; McKelvey et a l., 1980; Botsford et al., 1983; Thomas, 1985). In this study, we examine the relationship between advective transport of larvae and subsequent C. magister year-class strength along the southern Washington coast. We use data from trawl surveys of recently settled crab to measure year-class strength, thus minimizing potentially confounding effects associated with long time-lags (as required with commercial landings data). Aly», for the first time, we correct calculated (Bakun) wind data to account for wind veering in nearshore coastal waters. Our working hypothesis is that C. magister larvae do not reach preferred juvenile habitat, and that strong year classes are not possible unless a period of sustained landward transport occurs during the pelagic larval phase. Data and methods Study area Trawl sampling occurred along the southern Washington coast and consisted of both nearshore and estuarine components (Fig. 1). The nearshore area extended from the shoreline seaward to a depth of 59 m. This area encompasses nearly ha and constitutes approximately 40% of the suggested range of the southern Washington-northern Oregon crab stock (Barry, 1985). Also sampled were the two major estuaries along the Pacific coast of Washington, Grays Harbor (~8ha) and Willapa Bay (~11200ha). These estuaries have been described as important nursery areas for C. magister (Stevens and Armstrong, 1984; Armstrong and Gunderson, 1985; Gunderson et a l., 1990).

3 ICES mar. Sei. Symp., 199 (1995) Dungeness crab recruitment variability and Ekman transport o f larvae 169 Survey design and sampling protocol Generally, surveys were conducted throughout the study area during the months May through September, (Willapa Bay: ; see Gunderson et al. (1985) and Gunderson et al. (1990) for details). The nearshore area was sampled along fixed transects oriented east-west (Fig. 1) with trawl stations located at discrete depths, ranging from the edge of the surf zone at 5 m (3 fm) to the outer limit of crab abundance at 55 m (30 fm). Three depth strata grouped the two shallowest, the three intermediate, and the two deepest stations together. Stratum boundaries followed the mean lower low water, 14m (7.5fm), 41m (22.5fm), and 59m (32.5 fm) contours. Stratified random sampling occurred in the two estuaries (Fig. 1). Within each stratum, stations were randomly selected from a 1 km2 grid, with the constraint that no two stations were immediately adjacent to one another. Survey samples were collected in all three areas using a modified 3 m plumb staff beam trawl (Gunderson and Ellis, 1986). This system was scaled for use aboard a 6.4 m Boston Whaler (150 hp) within estuaries and for larger commercial fishing vessels (11 to 21 m) along the open coast. Effective width of the net was 2.3 m, while the estimated vertical opening was 0.6 m. Tows were taken parallel to isobaths in the nearshore (mean distance: 750 m) and, in the estuaries, along the main axis of the waterway (mean distance : 260 m). Crab present in trawl samples were sexed, counted and carapace widths measured inside the tenth anterolateral spines. Biological data Settlement success is measured here by the total abundance of 0+ (i.e., years since hatching) crab in the nearshore area during August of each year. The size range of 0+ crab was determined by inspection of modes in length-frequency plots, a relatively straightforward task for this age class (Fig. 2). Population estimates for each area were generated using area-swept procedures. However, because Willapa Bay was not sampled during the first two years of the survey ( ) and because the majority of new C. magister recruits occurred in the nearshore area ( mean: 94%; Fig. 3), the nearshore component was deemed the most appropriate index of abundance for this study. For simplicity, the annual index is discussed here as a Relative Abundance Factor (RAF), which is the ratio of abundance during the year in question to that of 1983, the period of lowest abundance. Physical data Alongshore windstress is an important forcing mechanism for large-scale coastal current fluctuations in the 4 3 c CD y 2 CD CL Carapace Width (cm) Figure 2. Representative C. magister length-frequency diagram from the 1985 trawl survey along the southern Washington coast. Note the clear distinction between 0+ (to the left of arrow) and subsequent age classes. Size limits in August of each year were the basis for calculations of 0+ crab population size by the area swept method. Pacific Northwest (Hickey, 1979, 1989; Halliwell and Allen, 1984). Calculated winds, as well as northward and eastward components of associated Ekman transport (ET), were obtained for a position central to the study area (46 55 N, 'W; Fig. 1) from the National Marine Fisheries Service Southwest Fisheries Center (D. Husby, Monterey, California). These wind vectors and associated transport measures (Bakun, 1973) are derived from 6-h pressure fields prepared by 0+ Abundance (millions) Nearshore Grays Harbor Willapa Bay Figure 3. Estimated abundance of 0+ C. magister in August ( ) in the three areas sampled by the trawl survey. Note that Willapa Bay was not sampled in 1983 and 1984 and that a large percentage of the total population routinely occurs in the nearshore area.

4 170 R. A. McConnaughey, D. A. Armstrong, and B. M. Hickey ICES mar. Sei. Symp., 199 (1995) the US Navy (Fleet Numerical Oceanographic Center). These data form an attractive set for analysis because they are temporally complete and derived by a common process and, as such, are used extensively as indicators of wind-driven currents. Ekman transport describes wind-driven circulation as a function of the wind stress (Sverdrup et al., 1942; Stacey et al., 1986). The ET data we obtained were resolved into northward and eastward vectors of depthintegrated ( mass ) transport per unit width of ocean surface (ts_1 km-1). In this form, ET is oriented 90 to the right (in the northern hemisphere) of the direction toward which the wind is blowing (Bakun, 1973), in keeping with Ekman s (1905) theory. We, however, have limited the ET rotation to 45, given the progressive deflection of wind-driven flow with depth (the familiar Ekman spiral ) and the near-surface orientation of C. magister zoeae and megalopae (Lough, 1976; Jacoby, 1982; Reilly, 1983; Jamieson and Phillips, 1988; Jamieson et al., 1989). An additional correction to the calculated winds and ET vectors was required so as to account for wind veering in coastal areas. In order to compensate for frictional effects, Bakun s (1973) estimate of surface winds incorporates a simple correction whereby wind speed is reduced by 30% and wind vectors rotated 15 counterclockwise. However, Halliwell and Allen (1984, 1987) demonstrated a misalignment between calculated and measured winds along the west coast of North America and concluded that Bakun s rotational correction is insufficient for coastal areas (see also Hsueh and Romea, 1983; Thomson, 1983). This discrepancy reflects a tendency for offshore w inds to veer parallel to the coastline in response to coastal topography, surface horizontal temperature gradients and diurnal winds. A supplemental correction accounts for this phenomenon and varies as a function of position along the coast, ranging from 5 to 50 counterclockwise over alongshore distances of a few hundred kilometers (Halliwell and Allen, 1987). Corrections which have been applied while computing corrected Ekman transport vectors are summarized in Figure 4. (a) Bakun data (initial condition) (b) Correction for nearsurface Ekman transport Ekman Wind (c) Correction to calculated (Bakun) winds Figure 4. Rotational corrections applied to (a) the original Bakun wind records to account for (b) the surface orientation of C. magister larvae (Ekman rotation limited to 45 clockwise) and (c) the misalignment between calculated and measured winds in coastal areas (original wind and associated Ekman transport vectors rotated 39 counterclockwise for our study area). Net correction (b + c) = 84 counterclockwise, which applies to C. magister larvae in coastal waters off southern Washington. Nearshore 0+ Abundance (millions) Results Crab settlement Settlement in the nearshore study area varied considerably over the 6-year period (Fig. 5). The strongest settlement occurred during 1985 (RAF = 44.9) and 1988 (RA F = 19.4). A moderately strong year class was recorded in 1984 (RAF = 9.3), while settlement was noticeably weaker during 1987 (RAF = 4.9), 1986 (RAF = 4.5), and 1983 (RAF = 1.0). Despite this variability, the nearshore was consistently the dominant area with Figure 5. August estimates of 0+ C. magister abundance in the nearshore area, Numbers above individual bars are relative abundance factors (RAF) and reference each estimate to the year of lowest abundance, 1983.

5 ICES mar. Sei. Symp., 199 (1995) (a) E o CL w c Ê c(0 E LU 1, Dungeness crab recruitment variability and Ekm an transport o f larvae , ; - 87 / / 84 / ^, 8 8 (b) CO JZ Eastward Ekman Transport (km) Figure 6. (a) Progressive vector diagrams of Ekman transport for the January to May C. magister larval period, Subplots are projected on a common coordinate system to facilitate comparisons and cumulative units of ts km have been omitted for clarity. Squares designate 30-day intervals between the endpoints, (b) Resultant vectors obtained by connecting the origin of each progressive vector diagram with the head of the final vector. Scale has been enlarged to show detail and years are as indicated. regard to 0+ abundance (98%, 98%, 84%, and 94% of total for ). Ekman transport Inspection of the progressive vector diagrams of ET for January-May, , suggests two main tendencies for the southern Washington coast (Fig. 6). Most notable is the near absence of sustained westward (seaward) transport during all years. With the exception of a relatively brief period during 1985 (January-early February) and another during 1988 (January), net nearsurface transport was consistently eastward (landward). Secondly, there was considerable interannual variability in both landward and northward ET. Strong northward and relatively weak landward transport characterized the years 1983 and In contrast, northward transport was considerably reduced during 1985 and 1988, while landward transport was substantial; significant northward transport ceased early in the larval period in both cases. The conditions during 1984 and 1987 were intermediate: ET during 1984 (to a greater degree) and 1987 (to a lesser degree), although northward, was moderated by a persistent landward component. Overall, the net direction of Ekman transport ranged 36, from a bearing of 31 in 1983 to 67 in 1985 (Fig. 6). Discussion The planktonic nature of C. magister larvae, coupled with rather restrictive juvenile habitat requirements, has prompted numerous investigations into the relationship between transport phenomena and ensuing year-class strength. These attempts have been largely unsuccessful due, at least in part, to shortcomings associated with use of commercial fishery landings as a proxy for year-class strength at settlement. Moreover, standard calculated winds have not been corrected (e.g., Johnson et a i, 1986), heretofore, so as to be appropriate for nearshore coastal waters. Using these new pieces of information, we are able to evaluate the prevailing recruitment paradigm for the species. The prevailing recruitment paradigm for C. m agister Based on field sampling off central California, it was hypothesized (Reilly, 1983; Hatfield, 1983) that C. magister zoeae are advected progressively seaward after hatching in nearshore waters and return to coastal areas as megalopae after considerable landward transport. This conclusion was based on patterns of spatial distribution for the five zoeal stages and intermolt stages of megalopae. However, the progressive seaward trans

6 R. A. McConnaughey, D. A. Armstrong, and B. M. Hickey ICES mar. Sei. Symp., 199 (1995) port (PST) hypothesis may simply be an artifact of very limited nearshore sampling effort and the correspondingly low numbers of late-stage larvae taken. For example, Reilly (1983) failed to catch stage V zoeae during four of the seven years studied while, in a similar yet smaller study off the Oregon coast, Lough (1976) found virtually no late stage larvae. Moreover, the unspecified mechanism is admittedly problematic given that the inferred direction of larval transport is opposed to the dominant cross-shelf circulation during winter (Hickey, 1979; Strub et al., 1987). We find no evidence in the ET series (Fig. 6) of sufficiently sustained seaward transport to account for quantities of late stage zoeae (age 2-3 months) presumed to be well offshore. Crossshelf movement of megalopae, on the other hand, is more tenable. The progressive vector diagrams of ET clearly indicate that surface waters may move landward during the period megalopae are present in the water column (Fig. 6) and Hatfield s (1983) evidence is rather compelling. However, it has been implied that offshore areas constitute the primary source of new recruits (Hatfield, 1983; Jamieson and Phillips, 1988; Jamieson e/a/., 1989), yet the origin (and relative contribution) of these megalopae is unknown and, as such, these studies, at best, provide only circumstantial evidence in support of the PST hypothesis. adult habitat. Although this idea contrasts with the notion of progressive seaward transport through ontogeny, it is nevertheless the only reasonable interpretation of our data. Similarly, Lough (1976), who found early larval stages close to the coast and some later stages further offshore, concluded that most C. magister larvae are retained nearshore by strong alongshore and onshore components of surface circulation. Because of restrictive juvenile habitat requirements, retention of larvae nearshore is conducive to successful settlement, given an outer depth limit of m beyond which new recruits were rarely encountered during six years of trawl sampling (including years of heavy settlement; Fig. 7). Median CPUE (n/ha) Dynamics of C. magister larvae along the southern Washington coast Our analyses suggest that C. magister larvae are retained nearshore during their ontogeny and that the strength of ensuing year classes of juveniles may be (inversely) related to the magnitude of alongshore transport during this period. C. magister settlement along the southern Washington coast was heaviest during 1985 (RAF = 44.9) and 1988 (RAF = 19.4; Fig. 5), while net ET during these years was landward with a relatively minor northward component (Fig. 6). The lowest levels of settlement during the six years studied occurred in 1983 and 1986 (RAF = 1.0 and 4.5) and ET during January- May of these years was strongly northward with a considerably reduced landward component (Fig. 6). Low recruitment during these years despite periods of strong landward flow during January suggests that the timing of landward flow as well as the net magnitude is important. Intermediate levels of settlement (1984: RAF = 9.3; 1987: RA F = 4.9) were associated with intermediate levels of landward and northward ET (Fig. 6). Rather persistent landward transport of near-surface waters during January-M ay is a characteristic feature of oceanic circulation over the Pacific Northwest shelf (Hickey, 1979, 1989; Fig. 6). Onshore flow will tend to minimize cross-shelf advective losses while retaining larvae in productive coastal waters near suitable juvenile/ Depth (meters) Figure 7. Median density (n/ha) of 0+ C. magister calculated using nearshore survey data for all months and all years. Note apparent outer depth limit between 37 and 46 m depth beyond which juvenile crab abundance (and presumably settlement by megalopae) is extremely low. Trawl sampling at 64 m (35 fm) occurred only during 1983 and was subsequently discontinued because of zero catches. Along the southern Washington coast, this outer depth limit corresponds to a distance of approximately 15 km from shore. Therefore, retention of larvae nearshore may be the product of a reproductive strategy (and surface-oriented larval swimming) which has evolved in response to time-averaged patterns in nearshore advective processes. The prevalence of winter and early spring spawning by Pacific demersal fish populations (Parrish et al., 1981) and other decapods (Rothlisberg and Miller, 1982; Jamieson and Phillips, 1988) suggests that significant adaptive value is attached to this strategy. Our conclusion concerning nearshore retention does not preclude cross-shelf input of recruits. Our evidence does suggest, however, that they are not of primary importance to C. magister populations along the coast of Washington. Alongshore current velocities can be considerable, yet this component of the coastal circulation has received scant attention with regard to C. magister larval advec-

7 ICES mar. Sei. Symp (1995) Dungeness crab recruitment variability and Ekm an transport o f larvae 173 tion. Indeed, the apparent absence of genetically distinct stocks (Soulé and Tasto, 1983), as well as synchronous patterns in commercial landings along the US west coast (Methot, 1986), suggest substantial alongshore linkages among (sub-)populations. Poor recruitment along the southern Washington coast during years of persistent wind-driven northward transport (e.g., 1983,1986; Fig. 6) implies that significant numbers of C. magister larvae were exported to the north. It is unclear, however, whether this represents a source of recruits for more northern populations or simply constitutes larval wastage. It is also unclear whether Washington receives recruits originating elsewhere. Jamieson et al. (1989) suggest that strong recruitment along the Washington coast in 1985 (Fig. 5) was due, at least in part, to an influx of larvae from the Vancouver Island stock; a reciprocal exchange may have occurred during 1986 when larvae exposed to sustained northward transport off Washington (Fig. 6) could have been spun north toward Vancouver Island. Furthermore, it is likely that Washington populations receive inputs of new recruits from southern populations, given the characteristic northward flow along the Pacific Northwest coast. Conclusions This was the first in a series of attempts by the authors to understand the recruitment dynamics of the commercially important Dungeness crab. In retrospect, several important factors were not considered. One of these was the geostrophic component of the alongshore circulation. In the Pacific Northwest, alongshore windstress can drive a relative (geostrophic) current that flows along the shelf. These currents functionally dominate alongshore flow and velocities can be considerable, particularly during winter. Also not addressed in this preliminary study was the functional basis for the outer depth limit of successful crab settlement. Identifying the responsible factor(s) could provide valuable insight into the spatial distribution of suitable settlement areas. These issues were subsequently addressed in McConnaughey et al. (1992,1993). As with all new hypotheses, certain cautions are in order until such time as various assumptions can be adequately tested. The spatial aspects of spawning are not well known. We have relied on the generally held assumption that spawning occurs close to shore. This is in keeping with female tagging studies (Diamond and Hankin, 1985; Hankin eta l., 1989), the relatively shallow water nature of the commercial fishery, and the prevalence of early stage zoeae close to shore (Lough, 1976; Reilly, 1983). Also, a well-defined spawnerrecruit relationship for crustaceans has not been described (Caddy, 1986) and information concerning C. magister spawner abundance from the male-only fisheries data is lacking. As such, no inference has been made here regarding interannual variability in magnitude of spawning stock. Evidence concerning the vertical distribution of C. magister larvae in the water column is scant. All available evidence suggests that brachyuran larvae in general (Sulkin, 1984) and C. magister larvae (Jacoby, 1982; Lough, 1976; Reilly, 1983), in particular, reside in near-surface waters where they are subject to winddriven circulation of the type considered here. Acknowledgements This research program was supported by an institutional grant from the Washington Sea Grant Program (No. NA86AA-D-SG044, Project R/F-68) and the US Army Corps of Engineers (No. DACW67-85-C-0033). The authors gratefully acknowledge the efforts of the numerous helpers in the field and the two anonymous reviewers who provided insightful comments. References Armstrong, D. A., and Gunderson, D. R The role of estuaries in Dungeness crab early life history: a case study in Grays Harbor, Washington. University of Alaska Sea Grant R ep., 85-3: Bakun, A Coastal upwelling indices, west coast of North America, N O A A Tech. Rep. NMFS-SSRF-671, Nat. Mar. Fish. Serv., Seattle. 103 pp. Barry, S Overview of the Washington coastal Dungeness crab fishery. University of Alaska Sea Grant Rep., 85-3: Botsford, L. W Effect of individual growth rates on expected behavior of the northern California Dungeness crab (Cancer magister) fishery. Can. J. Fish, aquat. Sei., 41: Botsford, L. W., Armstrong, D. A., and Shenker, J. M Oceanographic influences on the dynamics of commercially fished populations. In Coastal oceanography of Washington and Oregon, pp Ed. by M. R. Landry and B. M. Hickey. Elsevier Oceanography Series, Amsterdam. 607 pp. Botsford, L. W., Methot, R. D., Jr., and Johnston, W. E Effort dynamics of the northern California Dungeness crab (Cancer magister) fishery. Can. J. Fish, aquat. Sei., 40: Botsford, L. W., and Wickham, D. E Correlation of upwelling index and Dungeness crab catch. US Dept. Comm., Fish. Bull., 73: Botsford, L. W., and Wickham, D. E Behavior of agespecific, density dependent models and the northern California Dungeness crab (Cancer magister) fishery. J. Fish. Res. Bd Can. 35: Caddy, J. F Modeling stock-recruitm ent processes in Crustacea: some practical and theoretical perspectives. Can. J. Fish, aquat. Sei., 43: Cleaver, F. C Preliminary results of the coastal crab (Cancer magister) investigation. Wash. Dep. Fish. Biol. R ep., 49A: Diamond, N., and Hankin, D. G Movements of adult female Dungeness crabs (Cancer magister) based on tag recoveries. Can. J. Fish, aquat. Sei., 43: Ekman, V. 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8 174 R. A. McConnaughey, D. A. Armstrong, and B. M. Hickey ICES mar. Sei. Symp., 199 (1995) Gunderson, D. R., Armstrong, D. A., and Rogers, C Sampling design and methodology for juvenile Dungeness crab surveys. University of Alaska Sea Grant Rep., 85-3: Gunderson, D. R., Armstrong, D. A., Shi, Y., and McConnaughey, R. A Patterns of estuarine use by juvenile English sole (Parophrys vetulus) and Dungeness crab (Cancer magister). Estuaries, 13: Gunderson, D. R., and Ellis, I. E Development of a plumb staff beam trawl for sampling demersal fauna. Fish. Res., 4: Halliwell, G. R., Jr., and Allen, J. S Large-scale sea level response to atmospheric forcing along the west coast of North America, summer J. Phys. Oceanogr., 14: Halliwell, G. R., Jr., and Allen, J. S The large-scale coastal wind field along the west coast of North America, J. Geophys. Res., 92C: Hankin, D. G Proposed explanations for fluctuations in abundance of Dungeness crabs: a review and critique. University of Alaska Sea Grant Rep., 85-3: Hankin, D. G., Diamond, N., Mohr, M. S., and Ianelli, J Growth and reproductive dynamics of adult female Dungeness crabs (C ancermagister) in northern California. J. Cons. int. Explor. Mer, 46: Hatfield, S. E Intermolt staging and distribution of Dungeness crab, Cancer magister, megalopae. Calif. Dept. Fish Game Fish Bull., 172: Hickey, B. M The California Current system: hypotheses and facts. Prog. Oceanogr., 8: Hickey, B. M Patterns and processes of circulation over the Washington continental shelf and slope. In Coastal oceanography of Washington and Oregon, pp Ed. by M. R. Landry and B. M. Hickey. Elsevier Oceanography Series, Amsterdam. 607 pp. Hsueh, Y., and Romea, R. D A comparison of observed and calculated wintertime surface winds over the East China Sea. J. Geophys. Res., 88: Jacoby, C. A Behavioral responses of the larvae of Cancer magister Dana (1852) to light, pressure and gravity. Mar. Behav. Physiol., 8: Jamieson, G. S., and Phillips, A. C Occurrence of Cancer crab (C. magister and C. oregonensis) megalopae off the west coast of Vancouver Island, British Columbia. US Dept. Comm., Fish. Bull., 86: Jamieson, G. S., Phillips, A. C., and Huggett, W. S Effects of ocean variability on the abundance of Dungeness crab (Cancer magister) larvae. Can. Spec. Publ. Fish, aquat. Sei., 108: Johnson, D. F., Botsford, L. W., Methot, R. D., Jr., and Wainwright, T. C Wind stress and cycles in Dungeness crab (Cancer magister) catch off California, Oregon and Washington. Can. J. Fish, aquat. Sei., 43: Lough, R. G Larval dynamics of the Dungeness crab, Cancer magister, off the central Oregon coast, US Dept. Comm., Fish. Bull., 74: Love, M. S., and Westphal, W. V A correlation between annual catches of Dungeness crab, Cancer magister, and mean annual sunspot number. US Dept. Com m., Fish. Bull., 79: McConnaughey, R. A., Armstrong, D. A., Hickey, B. M., and Gunderson, D. R Juvenile Dungeness crab (Cancer magister) recruitment variability and oceanic' transport during the pelagic larval phase. Can. J. Fish, aquat. Sei., 49: McConnaughey, R. A., Armstrong, D. A., Hickey, B. M., and Gunderson, D. R Interannual variability in coastal Washington Dungeness crab (Cancer magister) populations: larval advection and the coastal landing strip. Fish. Oceanogr., 3: McKelvey, R., Hankin, D., Yanosko, K., and Snygg, C Stable cycles in multistage recruitment models: an application to the northern California Dungeness crab (Cancer magister) fishery. Can. J. Fish, aquat. Sei., 37: M ethot, R. D., Jr Management of Dungeness crab fisheries. Can. Spec. Publ. Fish, aquat. Sei., 92: Orensanz, J. M., and Gallucci, V. F A comparative study of postlarval life history schedules in four sympatric Cancer species. J. Crustacean Biol., 8: Parrish, R. H., Nelson, C. S., and Bakun, A Transport mechanisms and reproductive success of fishes in the California Current. Biol. 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Fish Game Fish Bull., 172: Stacey M. W., Pond, S., and LeBlond, P. H A windforced Ekman spiral as a good statistical fit to low-frequency currents in a coastal strait. Science, 233: Stevens, B. G., and Armstrong, D. A Distribution, abundance and growth of juvenile Dungeness crabs, Cancer magister, in Grays H arbor estuary, Washington. US Dept. Comm., Fish. Bull., 82: Strub P. T., Allen, J. S., and Smith, R. L Seasonal cycles of currents, temperatures, winds and sea level over the northeast Pacific continental shelf: 35 N to 48 N. J. Phys. Oceanogr., 92C: Sulkin, S. D Behavioral basis of depth regulation in the larvae of brachyuran crabs. Mar. Ecol. Prog. Ser., 15: Sverdrup, H. U., Johnson, M. W., and Fleming, R. H The oceans: their physics, chemistry and general biology. Prentice-Hall, New York pp. Thomas, D. H A possible link between coho (silver) salmon enhancement and a decline in central California Dungeness crab abundance. US Dept. Comm., Fish. Bull., 83: Thomson, R. E A comparison between computed and measured oceanic winds near the British Columbia coast. J. Geophys. Res., 88: Wild, P. W Effects of seawater tem perature on spawning, egg development, hatching success and population fluctuations of the Dungeness crab, Cancer magister. Calif. Coop. Oceanic Fish. Invest. R ep., 21: Wild, P. W The influence of seawater temperature on spawning, egg development and hatching success of the D ungeness crab, Cancer magister. Calif. Dept. Fish Game Fish Bull., 172:

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