ARTICLE IN PRESS

Deep-Sea Research II 52 (2005) 5172 www.elsevier.com/locate/dsr2

Abundance and distribution of large medusae in surface waters of the northern California Current
Cynthia L. Suchman,1, Richard D. Brodeur
Northwest Fisheries Science Center, National Marine Fisheries Service, Hateld Marine Science Center, Newport, OR 97365, USA Accepted 26 September 2004 Available online 7 January 2005

Abstract We used surface trawls to quantify abundance and distribution of four large medusaeChrysaora fuscescens, Aurelia labiata, Phacellophora camtschatica, and Aequorea sp.in the northern California Current, a coastal upwelling area, during June and August 2000 and 2002. C. fuscescens was most abundant, with a maximum of 77 per 1000 m3 (or 64 mg C/m3). Densities of A. labiata reached 10 per 1000 m3 (26 mg C/m3). Although medusae were widespread throughout the study region, a large proportion of the total catch of each species tended to be concentrated at relatively few stations. When we analyzed how the variability in distribution of medusae related to a suite of environmental parameters, latitude and either distance from shore or water depth proved important during each cruise. Analysis by species suggested habitat partitioning by C. fuscescens and A. labiata: C. fuscescens was more likely to be caught in nearshore shelf waters north of Cape Blanco, compared to A. labiata, which was more prevalent in south of Cape Blanco. These biological patterns were likely the result of a combination of physical processes, including advection and aggregation within regions of weak surface ow, coordinated behavior such as vertical migration, and location of benthic polyp stages. Studies such as this one provide important baseline information for ecosystem models and can be used to help evaluate impacts of jellysh blooms in coastal upwelling regions. r 2004 Elsevier Ltd. All rights reserved.

1. Introduction Many studies have demonstrated that gelatinous zooplankton can play a critical role in structuring coastal and estuarine ecosystems. These taxa can
Corresponding author. Fax: +1 434 982 3694.

E-mail address: csuchman@virginia.edu (C.L. Suchman). Current address: Virginia Graduate Marine Science Consortium, University of Virginia, Charlottesville, VA 22903, USA.
1

exert signicant predation pressure, to the point of initiating trophic cascades (e.g., Kideys, 2002). For example, inverse relationships between abundances of scyphomedusae (Cnidaria: Scyphozoa) and zooplankton or ichthyoplankton have been documented in various locations (Lindahl and Henroth, 1983; Moller, 1984; Behrends and Schneider, 1995; Omori et al., 1995). Moreover, shifts in climate, eutrophication, new introductions, and overshing may lead to changes in

0967-0645/$ - see front matter r 2004 Elsevier Ltd. All rights reserved. doi:10.1016/j.dsr2.2004.09.017

ARTICLE IN PRESS
52 C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172

populations of gelatinous zooplankton, which in turn may change predation on or competition with other components of the food web (Jackson et al., 2001; Mills, 2001; Parsons and Lalli, 2002). Recent research shows increase in biomass (Brodeur et al., 2002), shifts in distribution (Graham, 2001a), and unusually dense aggregations (Uye et al., 2003) of large medusae in various coastal environments. Because of a paucity of data, highly productive sheries, and the advective nature of the ecosystem, it is sometimes assumed that gelatinous zooplankton have not historically formed a major component of the food web in coastal upwelling areas (Mills, 2001; Parsons and Lalli, 2002). However, in the northern California Current, populations of large medusae appear seasonally each year, with maxima occurring in late summer or early fall (Shenker, 1984). Four species of large medusae are common in surface waters off the coast of Oregonthree scyphomedusae (Aurelia labiata, Chrysaora fuscescens, and Phacellophora camtschatica) and one hydromedusa (Aequorea sp.). Aequorea spp. in the Pacic have been called Aequorea victoria or Aequorea aequorea or Aequorea coerulescensange, without clear consensus on whether these are indeed separate species (Wrobel and Mills, 1998). Therefore, we choose to refer to Aequorea sp. in this paper. Data quantifying large medusae in coastal upwelling regions remain sparse. One study has documented populations of scyphomedusae in the northern California Current from May through September 1981 (Shenker, 1984). A recent study in the Benguela Current used a combination of trawling and visual observations to determine distribution of medusae (Sparks et al., 2001), expanding on an earlier study in the same area that collected medusae using bongo tows (Fearon et al., 1992). Chrysaora spp. were concentrated inshore of the upwelling front in both systems (Shenker, 1984; Graham, 1994; Sparks et al., 2001). Aequorea spp. were observed in more offshore-oceanic water in the Benguela system (Sparks et al., 2001), but overlapped with distributions of C. fuscescens (inshore) in the northern California Current (Shenker, 1984). Data on distributions of P. camtschatica and Aurelia spp. in upwelling regions are too limited to summarize.

Coastal upwelling areas also provide a good environment in which to investigate links between physical forcing and biological processes. The California Current, the eastern boundary current of the North Pacic subtropical gyre, ows to the south off the Oregon coast. Typically, northerly winds begin in late spring and continue intermittently through the summer months, causing Ekman transport of surface water offshore and upwelling of cold, nutrient-rich subsurface water near the coast (Huyer, 1983; Landry et al., 1989). Intense periods of high primary productivity follow wind-driven upwelling events and support high abundances of upper trophic levels. In addition, broader-scale climate effectsoperating at annual (e.g. El Nino) or decadal (Pacic Decadal Oscillation) time scales (Mantua et al., 1997; Hickey, 1998)mediate the magnitude of any seasonal signal and contribute to interannual variability. The study area includes two prominent geographic features that can modify the location of surface currents and recently upwelled water (Batchelder et al., 2002). Heceta Bank (44.21N, 124.91W) is a subsurface extension of the shelf associated with elevated primary productivity during summer months (Fig. 1) (Hickey, 1998). Water circulation on the bank is complex and may retain particles normally subject to offshore or downstream transport during upwelling (Oke et al., 2002; Barth et al., 2005). Cape Blanco (42.91N, 126.61W) is a coastal promontory, protruding further to the west than any other feature in an area with a relatively straight coastline (Fig. 1). North of Cape Blanco, upwelling is generally conned to nearshore, shelf waters. In the vicinity of Cape Blanco, however, southward-owing currents may be deected offshore (Barth et al., 2000), leading to meanders that transport cold, upwelled watercontaining associated planktonaway from the coast. As a result, composition and distribution of zooplankton communities can differ north and south of Cape Blanco (Peterson and Keister, 2002). In this study, we document the abundance and distribution patterns of large medusae in the northern California Current using data collected as part of the US Global Ocean Ecosystems

ARTICLE IN PRESS
C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172
45 N

53

1
Heceta Bank
44 N

Newport OR

Oregon
4 5 7
43 N

Cape Blanco 10
42 N

California

12
125 W

Crescent City, CA
124 W

Fig. 1. The study region off the west coast of the United States. The 200-m isobath is shown, marking the edge of the continental shelf. Mesoscale transects lines 1, 4, 5, 7, 10, and 12, are labeled, with location of diel studies in June (line 1) and August (line 4) 2002 marked with solid circles (d)

Dynamics (GLOBEC) program in the Northeast Pacic. The purpose of the GLOBEC program is to combine a variety of approaches to better understand the interaction among climate, physical processes, and biological productivity in coastal ecosystems (Batchelder et al., 2002). One component of the Northeast Pacic Program has involved mesoscale eld studies in the northern California Current (from Newport, Oregon to Crescent City, CA), with cruises completed in June and August of both 2000 and 2002. Anecdotal evidence and studies conducted two decades ago (Shenker, 1984; Pearcy et al., 1985) suggest that gelatinous zooplankton may be an importantalbeit understudiedcomponent of the food web in the northern California Current. We present new information on medusan habitat and population variability in this region. The study is important because very little quantitative data on large gelatinous organisms are available in coastal regions in general and upwelling areas in particular. These data are rare because they are difcult to collect with commonly used gear over sufcient spatial and temporal scales. They are critical, however, if we hope to form credible hypotheses about where and when population

maxima occur, predict how changes in other biological or physical parameters will affect medusan populations, and ultimately use population parameters to estimate the predation impact of these taxa on lower trophic levels.

2. Methods 2.1. Sampling We collected medusae during four cruises on chartered shing vessels as part of GLOBEC Northeast Pacic process studies in the California Current: 29 May11 June 2000 (hereafter referred to as June 2000), 29 July12 August 2000 (August 2000), 118 June 2002 (June 2002), and 118 August 2002 (August 2002). The study area was off the US West Coast between Newport, Oregon (44.61N, 124.01W) and Crescent City, California (41.71N, 124.11W) (Fig. 1). Each cruise began with a mesoscale survey of six onshore-offshore transect lines where stations were established 330 nautical miles (5.655.6 km) from shore. Once these transects were completed, sampling continued at ne-scale process stations,

ARTICLE IN PRESS
54 C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172

chosen based on biological or physical features of interest. Although the location of ne-scale stations varied among cruises, their distribution was similar, except during August 2000 when stations extended farther offshore. In June and August 2002, diel studies were performed to assess daynight differences in trawl catches. For 24 h, sampling was completed every 4 h at a single station (transect line 1, 58-m depth, on 78 June 2002; line 4, 116 m, on 78 August 2002) (Fig. 1). At each station, a number of biological and physical parameters were assessed (including temperature, salinity, chlorophyll a, neuston, medusae, and nekton). All sampling, except during the two diel studies in 2002, was completed during daylight. Replicate water samples to determine chlorophyll a were collected using a 2-L Niskin bottle deployed at a depth of 3 m. Water samples of 138 ml (shelf) or 281 ml (slope) were concentrated on GFF lters, with chlorophyll a measured in the laboratory using a Turner Designs 10-AU uorometer and standard uorometric techniques. Temperature and salinity proles to 100 m (or to 10 m shallower than the bottom depth) were obtained using a SeaBird SBE 19 CTD Proler. Contours of temperature (at 5 m) and chlorophyll a (at 3 m) over the study region were created from interpolation of station data using Surfer 7.0 software. Medusae and nekton were collected using a Nordic 264 rope trawl (30-m wide, 18-m deep) built by Nor-Eastern Trawl Systems, Inc., towed in surface waters for 30 min at 33.5 knots. Start and end latitude and longitude were recorded using Global Positioning System. Mesh size of the trawl ranged from 162.6 cm in the throat to 8.9 cm in the cod end, with a 6.1-m-long, 0.8-cm mesh liner sewn into the cod end. All medusae were counted and measured immediately after capture at sea. Each specimen was identied, each species was weighed in aggregate (wet weight), and individuals of each species were counted and bell diameters measured (to nearest mm). For exceptionally large hauls, a subsample (at least 50) of each species was measured, counted, and weighed, with the total number of medusae calculated based on total species weight for the haul.

2.2. Calculations of abundance and biomass Abundances of medusae (number per 1000 m3) were calculated by multiplying the effective mouth area of the trawl by the distance towed. Distance towed was determined using start and end latitude and longitude, corrected for the curvature of the earth. Effective mouth area of the trawl was estimated based on average medusa diameters and net mesh size (123 m2 for C. fuscescens, A. labiata, and P. camtschatica and 86 m2 for Aequorea sp.). Actual abundances were likely higher than those calculated for one or more of the following reasons: (1) large mesh size undersampled smaller individuals, (2) population maxima may have been below the depth sampled (18 m), (3) 30-min tows integrated over spatial scales larger than patches of medusae (leading to underestimates of maximum abundances), (4) nets may have clogged at stations were medusae were most aggregated, and (5) tows were not attempted at some inshore stations where medusae were particularly dense. Nevertheless, consistent trawling methods were employed during each cruise and therefore comparisons among tows, seasons, and years are appropriate. Carbon biomass (mg C/m3) for C. fuscescens and A. labiata at selected stations was determined by multiplying abundance by weight conversion (from count and wet weight measurements at sea). For C. fuscescens, carbon was calculated as 0.280% wet weight (Shenker, 1985); for A. labiata, carbon was assumed to be 0.163% wet weight (from data for A. aurita; Larson, 1986). 2.3. Statistical analyses We used an ordination technique to determine to what extent distribution of the four species of medusae could be described using environmental variables (Clarke and Warwick, 2001). The variables analyzed were: water depth, distance from shore, latitude, temperature (at 5 and 15 m), salinity (at 5 and 15 m), chlorophyll a (at 3 m), and neuston biovolume (Reese et al., 2005; S. Pool, unpublished data). First, we removed stations where no medusae were collected (1230% of tows, see Table 1), log(X+1) transformed the

ARTICLE IN PRESS
C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172 55

remaining species abundance data, and collapsed the species-by-station data into a BrayCurtis similarity matrix. Next, distributions of the environmental variables that were highly skewed (water depth, distance from shore, chlorophyll a, and neuston biomass) were log(X+1) transformed. Finally, we used a Spearman Rank Correlation statistic (r) to compare the environmental and species similarity matrices and determine which parameters accounted for the most variability in the distribution of medusae among stations (BIOENV function in PRIMER-E; Version 5, Clarke and Gorley, 2001). Although we used ordination analysis to evaluate all taxa and environmental variables simultaneously, this method required removing stations
Table 1 Summary of medusa catch data (n trawls per cruise) June 2000 (n 84) Frequency of occurrence (%) Aequorea sp. Aurelia labiata Chrysaora fuscescens Phacellophora camtschatica Total Number of medusae caught Aequorea sp. Aurelia labiata Chrysaora fuscescens Phacellophora camtschatica Total Wet weight medusae caught (kg) Aequorea sp. Aurelia labiata Chrysaora fuscescens Phacellophora camtschatica Total Maximum number per 1000 m3 (% species catch) Aequorea sp. Aurelia labiata Chrysaora fuscescens Phacellophora camtschatica

where no medusae were present. To include zero data and compare each species with target environmental variables, we used more traditional statistical techniques. For example, to evaluate distributions of each species with respect to the shelf break and Cape Blanco, we used w2 (with Yates correction) to determine whether frequency of occurrence varied among geographic regions (north or south of Cape Blanco, or stations shallower or deeper than 200 m). We also used a Spearman Rank Correlation to test the relationship between each medusa species and station variables of interest (chlorophyll a, temperature, salinity). We used a MannWhitney Rank Sum test to compare day and night samples during the two diel

August 2000 (n 78)

June 2002 (n 104)

August 2002 (n 99)

56 36 33 1 70 2656 968 5353 1 8978 80 1094 856 1 2031

33 38 29 14 63 10,087 1787 22,414 27 34,315 101 2162 6724 29 9016

74 35 61 20 88 5712 245 2435 33 8425 228 353 584 89 1254

55 49 55 39 88 0 2366 3128 10,985 1455 17,934 189 4911 5383 1921 12,403

3.0 (23) 1.0 (39) 8.3 (64) 0.004 (100)

11.7 (29) 1.7 (32) 76.8 (69) 0.023 (33)

5.3 (24) 0.32 (51) 2.3 (36) 0.014 (16)

1.9 (22) 10.3 (75) 20.1 (35) 2.4 (59)

Frequency of occurrence reports the percentage of trawls catching each species (total reports the percent of trawls catching at least one species). Number of medusae caught was uncorrected for volume towed or number of trawls. Wet weight was calculated by multiplying number caught by average wet weight/count conversion for each cruise. We also compared maximum single-station density (and percent of species catch) for each cruise.

ARTICLE IN PRESS
56 C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172

studies in 2002 and KolmogorovSmirnov tests to compare medusan size distribution between seasons and transects during 2002.

3. Results 3.1. Environmental conditions Upwelling of cold, saline subsurface ocean water followed development of northerly winds during late spring through summer months in this study region. Wind data for 2000 show upwellingfavorable winds prior to both the June and August cruises (Barth et al., 2005). During each cruise, water temperature at 5-m depth showed a crossshelf transition from warmer water offshore to cold, upwelled water near shore (Fig. 2). Since upwelling delivers nutrients that facilitate primary productivity, areas with elevated chlorophyll a were associated with nearshore cold water (Fig. 3). During both years, the geographic extent of upwelling was larger in August than in June with a corresponding increase in chlorophyll a concentration (Figs. 2 and 3). Upwelling and primary productivity were not uniformly distributed across the shelf; Heceta Bank and Cape Blanco were areas of enhanced upwelling and elevated chlorophyll a. During 2002, an anomalous inux of subarctic water into the region (Wheeler et al., 2003) led to lower sea surface temperatures and higher concentrations of chlorophyll a compared with 2000. 3.2. Medusae Large medusae (the hydromedusa Aequorea sp. and scyphomedusae C. fuscescens, A. labiata, and P. camtschatica) were widely distributed throughout the study region, caught in 6388% of trawls (Table 1). C. fuscescens dominated numerically (up to 22,414 individuals per cruise) and in terms of biomass (up to 6700 kg per cruise), though Aequorea sp. were also numerous (up to 10,087) and biomass of A. labiata could be relatively high (up to 4911 kg). In general, P. camtschatica was the least abundant species within the study region; during August 2002, however, catch of P.

camtschatica was substantially greater than during previous cruises, with biomass exceeding that of Aequorea sp. Periodic sampling over 24 h at one station during June and August 2002 did not show any signicant difference between day and night catches of C. fuscescens and Aequorea sp. (Mann Whitney Rank Sum test, p40:05) (Table 2), yet revealed high variability among trawls. Temperature and chlorophyll a were also not signicantly different between day and night samples (MannWhitney Rank Sum test, p40:05). Biomass of medusae increased 4 to 10-fold from June to August each year as population size and size of individuals increased (Tables 1 and 3). In general, abundances of medusae were higher in August than June (Figs. 47). However, in 2002, Aequorea sp. were more numerous in June than August (Fig. 6). Median size of individuals increased between June and August in 2002 (KolmogorovSmirnov test, po0:05), with the exception of P. camtschatica (where increase was not signicant, probably because relatively few individuals were measured) (Table 3). When we examined medusa size across transect lines, we found that medusae were often larger to the south. Diameters of C. fuscescens were larger along more southern transects in August (between lines 1 and 4, 4 and 5, 5 and 7; KolmogorovSmirnov test, po0:05) (Fig. 8). Aequorea sp. had a similar pattern during both June and August (KolmogorovSmirnov test, po0:05; between all transect lines except between lines 4 and 5 in August) (Fig 9). Aequorea sp. also showed a bimodal distribution along at least one line (7), suggesting two cohorts or two distinct populations during spring and summer 2002. We caught the largest biomass of medusae during the August 2002 cruise (Table 1). Although the dominant species, C. fuscescens, was actually more abundant in 2000 (because of a particularly large catch at one nearshore station, Fig. 4), A. labiata and P. camtschatica were more abundant in 2002 (Figs. 5 and 7). Medusae were not evenly distributed across sampling stations (Figs. 47). Even though stations extended farther offshore in August 2000 than during other cruises, catch of medusae

ARTICLE IN PRESS
C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172 57

45 N

June 2000

August 2000

44 N

43 N

Cape Blanco

42 N

125 W
45 N

124 W

126 W

125 W

124 W

June 2002

August 2002

44 N

43 N

42 N

125 W

124 W

125 W

124 W

Fig. 2. Contours of temperature (1C) from 5-m depth during each of four cruises. Station locations are marked with small solid circles. Two hundred meter isobath is shown as lighter, thinner line.

appeared more broadly distributed in 2002 than in 2000 (Frequency of Occurrence, Table 1; Figs. 47). In addition, a large proportion of the total catch of each species was concentrated in relatively few nearshore stations. For example, a single station accounted for between 35%

and 69% of total catch of C. fuscescens, with proportion varying by cruise. The highest abundance of C. fuscescens was caught at the station closest to shore on transect line 5 during August 2000 (69% of catch): 77 per 1000 m3 or 65 mg C/m3. Maximum abundances during

ARTICLE IN PRESS
58 C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172

June 2000
45 N

August 2000

44 N

43 N

Cape Blanco

42 N

125 W

124 W

126 W

125 W

124 W

June 2002
45 N

August 2002

44 N

43 N

42 N

125 W

124 W

125 W

124 W

Fig. 3. Contours of chlorophyll a (mg/l) from 3-m depth during each of four cruises. Station locations are marked with small solid circles. Two hundred meter isobath is shown as lighter, thinner line.

August 2002 were a similar order of magnitude and also at nearshore stations (Table 1, Figs. 4 and 5)for both C. fuscescens (35% of species

catch, 20 per 1000 m3 or 28 mg C/m3) and A. labiata (75% of species catch, 10 per 1000 m3 or 26 mg C/m3).

ARTICLE IN PRESS
C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172 Table 2 Abundances of medusae caught during day vs. night trawls over 24-h (mean7standard deviation per 1000 m3) June 2002 Day (n 5) Chrysaora fuscescens Aequorea sp. All medusae Chlorophyll a (mg/l) Temperature (1C) 0.06270.046 0.2870.28 0.3470.030 3.170.9 9.670.4 Night (n 2) 0.1270.051 0.02270.071 0.1570.020 4.170.3 9.970.4 August 2002 Day (n 4) 0.003070.0030 0.04970.060 0.05370.063 21.775.0 10.970.2 Night (n 3) 0.001270.0017 0.03670.051 0.03970.051 24.571.4 10.970.1 59

June 2002 sampling was on transect line 1 and August 2002 on line 4 (see Fig. 1). Differences between day and night data were not statistically signicant (p40:05; MannWhitney Rank Sum test).

Table 3 Mean diameter (cm) of medusae caught in trawls in the study region in 2002 (7standard deviation) June (n) Aequorea sp. Aurelia labiata Chrysaora fuscescens Phacellophora camtschatica 7.671.9 (1302) 30.476.8 (120) 17.375.4 (798) 28.1714.0 (27) August (n) 10.875.1 (814) 31.477.6 (575) 21.675.9 (679) 36.2716.1 (76) KolmogorovSmirnov po0:05 po0:05 po0:05 ns

ns: not signicant. Differences between June and August size distributions were signicant for all species except P. camtschatica (KolmogorovSmirnov test, po0.05).

3.3. Relationships among medusae and environmental variables Correlations of environmental variables with species ordinations showed that latitude and either water depth or distance from shore were the variables that explained the ordination patterns of medusae during every cruise (Table 4). Neuston biomass was also important in 2000. Overall correlation values ranging from r 0:46 in August 2002 to r 0:23 in June 2000 (Table 4). To illustrate distribution patterns of individual species, we plotted cumulative proportion of catch (including zero data) against distance from shore and latitude (Figs. 10 and 11). To include zero data in statistical analysis, we separated occurrence of each species into geographic habitats, bounded by either the 200-m isobath (characterized as on-shelf vs. offshore) or 42.81N (north or south of Cape Blanco). Relative to the proportion of trawls completed, C. fuscescens was concen-

trated on the shelf (o200-m depth) (w2 with Yates correction, po0:05 for all cruises but June 2000). Of the four taxa, C. fuscescens was consistently found closest to shore (Fig. 10). During all cruises but June 2000, A. labiata was caught to the south of Cape Blanco (w2 with Yates correction, po0:05) and south relative to sampling effort (Fig. 11). Patterns for Aequorea sp. and P. camtschatica were less consistent, although P. camtschatica often occupied more oceanic waters (August 2002) south of Cape Blanco (June and August 2002) (w2 with Yates correction, po0:05). Since C. fuscescens was most abundant close to shore throughout the summer (Figs. 4 and 10), we expected this species to inhabit primarily cold, saline water that had been subject to recent upwelling. Instead, C. fuscescens were present across a range of water temperatures and salinities sampled during all cruises except August 2000, which sampled stations with a wider range of temperatures than other cruises (Fig. 12). Therefore, C. fuscescens tolerated

ARTICLE IN PRESS
60 C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172

June 2000

August 2000

Chrysaora fuscescens (#/1000 m3) 44 0.0001 to 0.1 0.1 to 0.25 0.25 to 0.5 0.5 to 1 1 to 2.5 43 2.5 to 5 5 to 10 10 to 25 76 to 77
42
-125 -124

44

43

42 -125 -124

June 2002

August 2002

44

44

43

43

42 -125 -124

42 -125 -124

Fig. 4. Abundance and distribution of C. fuscescens during each of four cruises. Open circles represent size and location of catch in trawls (abundance reported as medusae per 1000 m3). Station locations are noted as small solid circles.

salinities from 30.1 to 34.0 psu and temperatures from 7.714.0 1C, and these variables proved relatively poor predictors of habitat for this medusa. Nevertheless, largest catches were concentrated in relatively cold, saline water, particularly in August (Fig. 12).

4. Discussion and conclusions 4.1. Patterns in abundance and distribution Large medusae in the study region were not randomly distributed. Moreover, even though

ARTICLE IN PRESS
C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172 61

June 2000

August 2000

Aurelia labiata (#/1000 m3) 0.0001 to 0.1 0.1 to 0.25 0.25 to 0.5 0.5 to 1 1 to 2.5 2.5 to 5 5 to 10 10 to 25

44

44

43

43

42 -125 -124

42 -125 -124

June 2002

August 2002

44

44

43

43

42 -125 -124

42 -125 -124

Fig. 5. Abundance and distribution of A. labiata during each of four cruises. Open circles represent size and location of catch in trawls (abundance reported as medusae per 1000 m3). Station locations are noted as small solid circles.

individuals of at least one species were caught in a majority of surface trawls deployed over four cruises, patterns varied based on species, season, and year (Figs. 10 and 11). For plankton populations, particularly in coastal upwelling regions such as the California Current, interactions between physics and biology may quickly lead to variation across time and space. In this study, we determined abundance of medusae concurrent

with other physical and biological parameters, providing more information about which factors may inuence the location and extent of patches of medusae. Undoubtedly, local density maxima occurred at ner scales than we sampled during 30-min tows. Our study was designed to explore mesoscale distribution; investigation of ner-scale processes requires other techniques or technology involving

ARTICLE IN PRESS
62 C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172

June 2000

August 2000

Aequorea sp. (#/1000 m3 ) 0.0001 to 0.1 0.1 to 0.25 0.25 to 0.5 0.5 to 1 1 to 2.5 2.5 to 5 5 to 10 10 to 25

44

44

43

43

42 -125 -124

42 -125 -124

June 2002

August 2002

44

44

43

43

42 -125 -124

42 -125 -124

Fig. 6. Abundance of Aequorea sp. during each of four cruises. Open circles represent size and location of catch in trawls (abundance reported as medusae per 1000 m3). Station locations are noted as small solid circles.

open-water diving or submersible camera systems (e.g., Graham et al., 2003). 4.1.1. Nearshore retention of Chrysaora fuscescens C. fuscescens, the most abundant of the four species caught in this study, was always most dense

at the most shoreward station sampled along each transect (Fig. 4). Because we were limited to water deep enough to deploy the trawl free of hazards such as shing gear and rocks, we were unable to assess the shoreward extent of the population, but C. fuscescens were abundant just beyond the jetty

ARTICLE IN PRESS
C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172 63

June 2000

August 2000

Phacellophora camtschatica (#/1000 m3 ) 0.0001 to 0.1 0.1 to 0.25 0.25 to 0.5 0.5 to 1 1 to 2.5

44

44

43

43

42 -125 -124

42 -125 -124

June 2002

August 2002

44

44

43

43

42 -125 -124

42 -125 -124

Fig. 7. Abundance and distribution of P. camtschatica during each of four cruises. Open circles represent size and location of catch in trawls (abundance reported as medusae per 1000 m3). Station locations are noted as small solid circles.

off Newport, OR, and often occurred in nearby Yaquina Bay during ood tide (E. Daly, Oregon State University, personal communication). Similar nearshore distributions of Chrysaora spp. have been noted before in upwelling areas, including off Oregon (Shenker, 1984), Monterey Bay, CA (Graham, 1994), and Namibia (Sparks et al.,

2001). Because ephyrae are released from benthic polyps, presumably attached to hard substrate in protected estuaries and embayments (Heitstuman, 1994) or under rocks protruding from reefs on the shelf, the planktonic part of the life cycle begins close to shore. However, coastal upwelling areas are highly advective, with bulk transport of surface

ARTICLE IN PRESS
64 C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172

0.6 0.5 0.4 0.3 0.2 0.1 0.0 0.6 0.5 0.4 0.3 0.2 0.1 0.0 0.6 0.5 0.4 0.3 0.2 0.1 0.0 0.6 0.5 0.4 0.3 0.2 0.1 0.0 0.6 0.5 0.4 0.3 0.2 0.1 0.0 0.6 0.5 0.4 0.3 0.2 0.1 0.0

1
June (15.5 4.5 cm, n=203) August (19.1 4.2 cm, n=90)

June (17.2 5.7 cm, n=117) August (17.5 5.1 cm, n=88)

proportion of medusae by transect

June (14.2 4.5 cm, n=22) August (24.2 5.6 cm, n=75)

June (14.9 4.0 cm, n=72) August (26.8 5.3 cm, n=74)

10

June (19.3 4.8 cm, n=84) August (30.7 5.3 cm, n=3)

12

June (20.3 5.1 cm, n=84)

05

35 30 -

510

25

25 -

30

bell diameter (cm)

Fig. 8. Diameter of C. fuscescens in 2002 by transect line (noted in upper left corner of each graph). See Fig. 1 for transect locations. Legend shows mean (cm)7standard deviation.

water in the California Current offshore and to the south during summer (Huyer, 1983). Although in some areas, fronts may be sufcient to retain populations of C. fuscescens, it is likely that in the northern California Current, both

20 -

10 -

15 -

physical and biological processes were important in determining nearshore distribution. Graham (1993, 1994) hypothesized that in Monterey Bay, C. fuscescens remain in a warm, low-chlorophyll shadow of water that persists shoreward of the

35 -

40

15

20

ARTICLE IN PRESS
C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172 65

0.5 0.4 0.3 0.2 0.1 0.0 0.5 0.4 0.3 0.2 0.1 0.0

1 June (7.2 1.8 cm, n=134) August (5.1 1.0 cm, n=13)

4 June (7.6 2.1 cm, n=145) August (9.4 4.1 cm, n=140)

proportion of medusae by transect

0.5 0.4 0.3 0.2 0.1 0.0 0.5 0.4 0.3 0.2 0.1 0.0 0.5 0.4 0.3 0.2 0.1 0.0 0.5 0.4 0.3 0.2 0.1 0.0

5 June (6.4 2.0cm, n=143) August (9.0 2.4 cm n=90)

7 June (7.1 1.7 cm, n=280) August (12.0 5.9 cm n=181)

10

June (7.7 2.1cm, n=150) August (18.1 4.4 cm, n=42)

12 June (8.9 1.5 cm, n=79)

0-5 5-7.5 .5-10 -12.5 .5-15 -17.5 .5-20 -22.5 .5-25 5-30 2 7 12 10 15 22 20 17

bell diameter (cm)

Fig. 9. Diameter of Aequorea sp. in 2002 by transect line (noted in upper left corner of each graph). See Fig. 1 for transect locations. Legend shows mean (cm)7standard deviation.

upwelling front. Similar features are not known to occur regularly along the Oregon coastline. Concurrent with our surface trawls during June and

August 2000, Barth et al. (2005) measured current velocities to 200 m along all transects using an Acoustic Doppler Current Proler (ADCP). They

ARTICLE IN PRESS
66 C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172

Table 4 Combinations of environmental and station variables that yield the best relationship between biotic and abiotic similarity matrices as measured by Spearmans rank correlation (r) Sample date June 2000 August 2000 June 2002 August 2002 Best variable combination Water depth, latitude, neuston biomass, distance from shore Latitude, distance from shore, neuston biomass Distance from shore, latitude Latitude, water depth Spearmans correlation 0.23 0.38 0.31 0.46

Variables describing distribution of the four species of medusae are listed in relative order of importance to the model.

June 2000

August 2000

1.0 0.8 0.6 0.4 cumulative proportion of catch 0.2 0.0 0 20 40 60 80 0 20 40 60 80 100 120 140

June 2002

August 2002

1.0 0.8 0.6 0.4 0.2 0.0 0 20 40 60 80 0 20 40 60 80 distance from shore (km)

Aequorea sp. Aurelia labiata Chrysaora fuscescens Phacellophora camtschatica sampling effort

Fig. 10. Cumulative catch of each medusa species compared with sampling effort by distance from shore. C. fuscescens was consistently caught very close to shore.

ARTICLE IN PRESS
C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172 67

June 2000

August 2000

1.0 0.8 0.6 0.4

cumulative proportion of catch

0.2 0.0 42 N
June 2002

43 N

44 N

42 N
August 2002

43 N

44 N

1.0 0.8 0.6 0.4 0.2 0.0 42 N 43 N 44 N 42 N 43 N 44 N latitude (South to North) Aequorea sp. Aurelia labiata Chrysaora fuscescens Phacellophora camtschatica sampling effort

Fig. 11. Cumulative catch of each medusa species compared with sampling effort by latitude. C. fuscescens was generally relatively north (particularly in August) and A. labiata and P. camtschatica to the south compared to station locations.

showed a complex and dynamic current structure in the upper 200 m of the water column, with areas of onshore and northward ow embedded in overall ow offshore and to the south (Pierce, 2004). Unlike mesoscale features associated with the upwelling jet that can persist for several weeks once formed, nearshore currents (o100-m water depth) are particularly responsive to shifts in wind direction, and can uctuate on the order of days (Barth et al., 2005). Like nearshore copepods in this system (Peterson et al., 1979), C. fuscescens may vary its vertical

location to move among water masses, using opposing currents to maintain horizontal position and resist advection. Chrysaora spp. have exhibited light-mediated vertical migrations (Schuyler and Sullivan, 1997; Brierley et al., 2001), and other scyphomedusae have used oriented swimming to maintain populations in various environments (Marshall and Hicks, 1962; Hamner et al., 1994). Although we did not see evidence of vertical migration during diel studies in 2002 (Table 2), our sampling integrated the upper 18 m of the water column. If shallow-amplitude migrations

ARTICLE IN PRESS
68 C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172

15 14 13 12 11

June 2000

temperature (C)

10 9 30 15 14 13 12 11 31 32 33

18 17 16 15 14 13 12 11 10 9 8

August 2000

32.0

32.5

33.0

33.5

34.0

June 2002
13 12 11 10 10 9 8 30 31 32 33 34 9 8

August 2002

>33% C. fuscescens catch >10% C. fuscescens catch C. fuscescens present C. fuscescens not present

temp. (C)

32.0 salinity (psu)

32.5

33.0 10 9 8

33.5

34.0

33.8 33.9 salinity (psu)

Fig. 12. Occurrence of C. fuscescens with respect to temperature and salinity measured at 5-m depth. Stations comprising relatively large catches are labeled with solid symbols.

occurred, they may not have been detected. Anecdotal evidence suggests that C. fuscescens in the northern California Current form dense layers that migrate between the surface at night to as deep as 30 m during daytime (C. Suchman, unpublished data; J. Burke, Oregon Coast Aquarium, personal communication). Because Heceta Bank (Fig. 1) served as a retention area for water masses (Oke et al., 2002; Barth et al., 2005), with elevated densities of phytoplankton (Fig. 3) and euphausiids (Ressler et al., 2005), we might have expected to see high abundances of C. fuscescens over the entire bank rather than merely at the most shoreward stations (Fig. 4). Instead, population maxima

of C. fuscescens were independent of mesoscale patterns in physics and biology on Heceta Bank, supporting the notion that this population resisted entrainment in the northern California Current. On the other hand, in 2002 we saw larger individuals along more southern transects (Fig. 8). If growth rates did not vary with latitude and polyps released ephyrae synchronously throughout the study area, the size pattern could have been caused by southward displacement of older medusae. Until data on individual growth rates are available and we know the source locations of polyps, this hypothesis will be impossible to address.

ARTICLE IN PRESS
C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172 69

4.1.2. Southern distribution of Aurelia labiata A. labiata was more likely to be collected south of Cape Blanco, particularly during August sampling periods (Figs. 5 and 11). In fact, latitude was the most salient factor related to medusan population structure during August 2000 and 2002 (Table 4). A more southern distribution would result if the source of the A. labiata population was also south of Cape Blanco and medusae exhibited behavior to enhance retention. Aurelia spp. polyps have been found in Yaquina Bay at Newport, OR (Heitstuman, 1994) and ephyrae have been collected during spring months at nearshore stations on transect line 1 (Fig. 1) (Suchman, unpublished data). A. labiata has been caught off the coast of Washington (Suchman, unpublished data), and large populations have been recorded in Prince William Sound (Purcell et al., 2000), so our study area was apparently not near the northern limit of its range. Although early stages of A. labiata are found across a broad geographic range, polyp distribution and ephyra production may be centered south of Cape Blanco. Alternatively, population maxima may have been found to the south because over the course of the summer, individuals from areas north of Cape Blanco were more likely subject to offshore advection. North-south separation of copepod communities has been documented in this region and attributed to the tendency for offshore meanders to develop south of Cape Blanco (Peterson and Keister, 2002). Interannual differences in location of A. labiata suggested that mesoscale physical features inuenced the distribution pattern of this species. In August 2000, maximum densities clustered around Cape Blanco (Fig. 5). These also corresponded to locally high densities of chlorophyll a (Fig. 3) and euphausiids (Ressler et al., 2005). A large anticyclonic meander in the coastal jet persisted just off Cape Blanco during this period (Barth et al., 2005; Pierce, 2004). Relatively weaker current velocities inshore of this feature might have facilitated aggregation of plankton across multiple trophic levels. In contrast, during August 2002, A. labiata was most abundant farther to the south of Cape Blanco (Fig. 5). Hydrographic and velocity data from August 2002 suggested that the southward-

owing jet was well offshore of the shelf break (J. Barth, Oregon State University, personal communication; Pierce, 2004), leading to a current structure less inuenced by Cape Blanco compared to 2000. Although A. labiata are known to form dense aggregations enhanced through swimming behavior (Hamner et al., 1994; Purcell et al., 2000), these data suggested thatlike phytoplankton and euphausiidslocation of population maxima may have been subject to modication based upon the physical environment. In addition, the southern distribution of A. labiata led to habitat partitioning with C. fuscescens. In 2002 in particular, although medusae were present in more trawls than in 2000 (Table 1), C. fuscescens was distributed north of Cape Blanco whereas A. labiata was more prevalent south of the cape (Fig. 11). Habitat partitioning among dominant species of medusae also has been reported in the northern Benguela Current (Sparks et al., 2001), and may be common along coastlines where persistent populations of medusae compete for prey resources. 4.2. Long-term changes in populations of medusae One motivation to estimate gelatinous zooplankton populations comes from a concern that these taxa may be increasing in abundance and expanding to new areas. Jellysh blooms, or unusual densities of these animals, can be caused either by a sustained increase in reproductive output, or physical aggregation events leading to apparent increases in population size (which may in fact only be a distributional shift) (Mills, 2001). Because this study only encompassed 2 years, we have insufcient data to address long-term changes in medusan populations in the region. However, another study conducted 20 years ago assessed large medusae in an area to the north of our work (focused off the Columbia River and Newport, OR) (Shenker, 1984). Medusae were collected using different gear (purse seine) with smaller mesh, but maximum biomass of C. fuscescens was comparable (50 mg C/m3 in 1981 vs. 64 mg C/m3 in 2000 and 28 mg C/m3 in 2002). Aggregation patternsnearshore population maximalikewise were similar. Moreover, biomass of

ARTICLE IN PRESS
70 C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172

gelatinous zooplankton caught during a triennial groundsh trawl survey off the US west coast since 1980 declined during El Nino years but otherwise showed no consistent trend (M. Wilkins, Alaska Fisheries Science Center, NOAA, personal communication). These limited data have provided no evidence that abundance or distribution of C. fuscescens has substantially changed over the past 20 years off the coast of Oregon. 4.3. Impact of medusae on the food web of the northern California Current How important are medusae to the ecosystem? To answer this question, we need to know not only how many medusae are present but also their feeding rates relative to zooplankton production rates. Maximum biomass of C. fuscescens and A. labiata reached 2664 mg C/m3, levels comparable to peak copepod biomass in the region over the past several years (Schwing et al., 2002). It is in these nearshore, high-density patches that predation rates may be sufcient to deplete the standing stock of zooplankton or ichthyoplankton. Although feeding rates by large medusae in the northern California Current are not yet known, both crustacean and gelatinous zooplankton should be vulnerable to predation (e.g., Strand and Hamner, 1988; Purcell and Sturdevant, 2001). In other coastal environments, large medusae ingested up to several thousand prey items per day (Graham, 2001b; Purcell, 2003). Moreover, medusae may inuence the food web in the northern California Current beyond simple ingestion of zooplankton. Indirect effects include competition with pelagic shes for similar prey (e.g., Purcell and Sturdevant, 2001). Not all interactions are detrimental, however. Gelatinous zooplankton provide food for upper trophic levels including sea turtles, seabirds, and several sh species (Purcell and Arai, 2001), and medusae may provide refuge habitat for juvenile sh (Brodeur, 1998). Given their seasonal and interannual persistence, areas of high biomass, interaction with various other components of the food web, and the potential for changes in populations through natural or human-mediated means, it is important to consider the effect of large medusae

on ecosystem dynamics in the northern California Current. Only through collection of long-term data and assessment of trophic interactions with respect to gelatinous zooplankton, will food web models in this region be complete.

Acknowledgements This research was supported by grants from NOAA through the US GLOBEC Program (contribution #481). C.L.S. was supported by a National Research Council Postdoctoral Research Associateship. We appreciate comments from J. Barth, H. Batchelder, R. Emmett, J. Keister, W. Pearcy, P. Ressler, J. Zamon, and two anonymous reviewers on earlier drafts of the manuscript. We thank the captains and crews of the FV Frosti and FV Sea Eagle as well as the following individuals who participated in research cruises and counted, weighed, and measured medusae: C. Anderson, B. Baldwin, P. Bentley, R. Emmett, J. Fisher, M. House, E. Locke, T. Miller, J. Popp Noskov, S. Pool, D. Reese, J. Richards, T. Sandell, and T. Wenham. S. Pool and J. Popp Noskov maintained the database and provided environmental data.

References
Barth, J.A., Pierce, S.D., Smith, R.L., 2000. A separating coastal upwelling jet at Cape Blanco, Oregon and its connection to the California Current System. Deep-Sea Research II 47, 783810. Barth, J.A., Pierce, S.D., Cowles, T.J., 2005. Mesoscale structure and its seasonal evolution in the northern California Current system. Deep-Sea Research II, this issue [doi:10.1016/j.dsr2.2004.09.026]. Batchelder, H.P., Barth, J.A., Kosro, P.M., Strub, P.T., Brodeur, R.D., Peterson, W.T., Tynan, C.T., Ohman, M.D., Botsford, L.W., Powell, T.M., Schwing, F.B., Ainley, D.G., Mackas, D.L., Hickey, B.M., Ramp, S.R., 2002. The GLBOEC Northeast Pacic California Current System Program. Oceanography 15, 3647. Behrends, G., Schneider, G., 1995. Impact of Aurelia aurita medusae (Cnidaria, Scyphozoa) on the standing stock and community composition of mesozooplankton in the Kiel Bight (western Baltic Sea). Marine Ecology Progress Series 127, 3945. Brierley, A.S., Axelsen, B.E., Buecher, E., Sparks, C., Boyer, H., Gibbons, M.J., 2001. Acoustic observations of jellysh

ARTICLE IN PRESS
C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172 in the Namibian Benguela. Marine Ecology Progress Series 210, 5566. Brodeur, R.D., 1998. In situ observations of the association between juvenile shes and scyphomedusae in the Bering Sea. Marine Ecology Progress Series 163, 1120. Brodeur, R.D., Sugisaki, H., Hunt, G., 2002. Increases in jellysh biomass in the Bering Sea: implications for the ecosystem. Marine Ecology Progress Series 233, 89103. Clarke, K.R., Gorley, R.N., 2001. PRIMER v5: User Manual/ Tutorial. Primer-E, Plymouth. Clarke, K.R., Warwick, R.M., 2001. Change in marine communities: an approach to statistical analysis and interpretation, second ed. Primer-E, Plymouth. Fearon, J.J., Boyd, A.J., Schulein, F.H., 1992. Views on the biomass and distribution of Chrysaora hysoscella (Linne, 1766) and Aequorea aequorea (Forskal, 1775) off Namibia 19821989. Scientia Marina 56, 7585. Graham, W.M., 1993. Spatio-temporal scale assessment of an upwelling shadow in northern Monterey Bay, California. Estuaries 16, 8391. Graham, W.M., 1994. The physical oceanography and ecology of upwelling shadows. Ph.D. Thesis, University of California, Santa Cruz, USA. Graham, W.M., 2001a. Numerical increases and distributional shifts of Chrysaora quinquecirrha (Desor) and Aurelia aurita (Linne) (Cnidaria: Scyphozoa) in the northern Gulf of Mexico. Hydrobiologia 451, 97111. Graham, W.M., 2001b. Size-based prey selectivity and dietary shifts in the jellysh Aurelia aurita. Journal of Plankton Research 23, 6774. Graham, W.M., Martin, D.L., Martin, J.C., 2003. In situ quanticiation and analysis of large jellysh using a novel video proler. Marine Ecology Progress Series 254, 129140. Hamner, W.M., Hamner, P.P., Strand, S.W., 1994. Sun compass migration by Aurelia aurita (Scyphozoa)population retention and reproduction in Saanich Inlet, British Columbia. Marine Biology 119, 347356. Heitstuman, T.M., 1994. Aspects of the biology and culture of scyphomedusae of the Oregon coast. M.S. Thesis, Oregon State University, Corvallis, Oregon, USA. Hickey, B.M., 1998. Coastal oceanography of western North America from the tip of Baja California to Vancouver Island. In: Robinson, A.R., Brink, K.H. (Eds.), The Sea, vol. 11. Wiley, New York, pp. 345393. Huyer, A., 1983. Coastal upwelling in the California Current System. Progress in Oceanography 12, 259284. Jackson, J.B.C., Kirby, M.X., Berger, W.H., Bjorndal, K.A., Botsford, L.W., Bourque, B.J., Bradbury, R.H., Cooke, R., Erlandson, J., Estes, J.A., Hughes, T.P., Kidwell, S., Lange, C.B., Lenihan, H.S., Pandol, J.M., Peterson, C.H., Steneck, R.S., Tegner, M.J., Warner, R.R., 2001. Historical overshing and the recent collapse of coastal ecosystems. Science 293, 629638. Kideys, A.E., 2002. Fall and rise of the Black Sea ecosystem. Science 297, 14821483. 71

Landry, M.R., Postel, J.R., Peterson, W.K., Newman, J., 1989. Broad-scale distributional patterns of hydrgraphic variables on the Washington/Oregon shelf. In: Landry, M.R., Hickey, B.M. (Eds.), Coastal Oceanography of Washington and Oregon. Elsevier Oceanography Series, vol. 46. Elsevier, New York, pp. 137. Larson, R.J., 1986. Water content, organic content, and carbon and nitrogen composition from medusae from the Northeast Pacic. Journal of Experimental Marine Biology and Ecology 99, 107120. Lindahl, O., Henroth, L., 1983. Phyto-zooplankton community in coastal waters of western Swedenan ecosystem off balance? Marine Ecology Progress Series 10, 119126. Mantua, N.J., Hare, S.R., Zhang, Y., Wallace, J.M., Francis, R.C., 1997. A Pacic interdecadal climate oscillation with impacts on salmon production. Bulletin of the American Meteorological Society 78, 10691079. Marshall, N., Hicks, S.D., 1962. Drift of medusae and their distribution in relation to the hydrography of the Niantic River, Connecticut. Limnology and Oceanography 7, 268269. Mills, C.E., 2001. Jellysh blooms: are populations increasing globally in response to changing ocean conditions? Hydrobiologia 451, 5568. Moller, H., 1984. Reduction of a larval herring population by jellysh predator. Science 224, 621622. Oke, P.R., Allen, R.N., Egbert, G.D., 2002. A modeling study of the three-dimensional continental shelf circulation off Oregon. Part II: dynamical analysis. Journal of Physical Oceanography 32, 13831403. Omori, M., Ishii, H., Fujinaga, A., 1995. Life history strategy of Aurelia aurita (Cnidaria, Scyphomedusae) and its impact on the zooplankton community of Tokyo Bay. ICES Journal of Marine Science 52, 597603. Parsons, T.R., Lalli, C.M., 2002. Jellysh population explosions: revisiting a hypothesis of possible causes. La mer 40, 111121. Pearcy, W., Fisher, J., Brodeur, R., Johnson, S., 1985. Effects of the 1983 El Nino on coastal nekton off Oregon and Washington. In: Wooster, W.S., Fluharty, D.L. (Eds.), El Nino North: Nino Effects in the Eastern Subarctic Pacic Ocean. Washington Sea Grant Program, University of Washington, Seattle. Peterson, W.T., Keister, J.E., 2002. The effect of a large cape on distribution patterns of coastal and oceanic copepods off Oregon and northern California during the 19981999 El NinoLa Nina. Progress in Oceanography 53, 389411. Peterson, W.T., Miller, C.B., Hutchinson, A., 1979. Zonation and maintenance of copepod populations in the Oregon upwelling zone. Deep-Sea Research 26A, 467494. Pierce, S.D., 2004. Mesoscale and nescale mapping of physical and biological elds in the northern California Current system: acoustic Doppler current proler data. www Page, http://damp.coas.oregonstate.edu/globec/nep/adcp. Purcell, J.E., 2003. Predation on zooplankton by large jellysh, Aurelia labiata, Cyanea capillata and Aequorea aequorea, in

ARTICLE IN PRESS
72 C.L. Suchman, R.D. Brodeur / Deep-Sea Research II 52 (2005) 5172 R.J., Mantyla, A.W., Ohman, M.D., Peterson, W.T., Smith, R.T., Sydeman, W.J., Venrick, E., Wheeler, P.A., 2002. The state of the California Current, 20012002: will the California Current system keep its cool, or is El Nino looming? CalCOFI Report 43, 3168. Shenker, J.M., 1984. Scyphomedusae in surface waters near the Oregon coast, MayAugust 1981. Estuarine, Coastal and Shelf Science 19, 619632. Shenker, J.M., 1985. Carbon content of the neritic scyphomedusa Chrysaora fuscescens. Journal of Plankton Research 7, 169173. Sparks, C., Buecher, E., Brierley, A.S., Axelsen, B.E., Boyer, H., Gibbons, M.J., 2001. Observations on the distribution and relative abundance of the scyphomedusan Chrysaora hysoscella (Linne, 1766) and the hydrozoan Aequorea aequorea (Forskal, 1775) in the northern Benguela ecosystem. Hydrobiologia 451, 275286. Strand, S.W., Hamner, W.M., 1988. Predatory behavior of Phacellophora camtschatica and size-selective predation upon Aurelia aurita (Scyphozoa: Cnidaria) in Saanich Inlet, British Columbia. Marine Biology 99, 409414. Uye, S., Fujii, N., Takeoka, H., 2003. Unusual aggregations of the scyphomedusa Aurelia aurita in coastal waters along western Shikoku, Japan. Plankton Biology and Ecology 50, 1721. Wheeler, P.A., Huyer, A., Fleischbein, J., 2003. Cold halocline, increased nutrients and higher chlorophyll off Oregon in 2002. Geophysical Research Letters 30, 8021. Wrobel, D., Mills, C., 1998. Pacic coast pelagic invertebrates: a guide to the common glelatinous animals. Sea Challengers and Monterey Bay Aquarium, Monterey, California.

Prince William Sound, Alaska. Marine Ecology Progress Series 246, 137152. Purcell, J.E., Arai, M.N., 2001. Interactions of pelagic cnidarians and ctenophores with sh: a review. Hydrobiologia 451, 2744. Purcell, J.E., Sturdevant, M.V., 2001. Prey selection and dietary overlap among zooplanktivorous jellysh and juveline shes in Prince William Sound, Alaska. Marine Ecology Progress Series 210, 6783. Purcell, J.E., Brown, E.D., Stokesbury, K.D.E., Haldorson, L.H., Shirley, T.C., 2000. Aggregations of the jellysh Aurelia labiata: abundance, distribution, association with age-0 walleye pollock, and behaviors promoting aggregation in Prince William Sound, Alaska USA. Marine Ecology Progress Series 195, 145158. Reese, D.C., Miller, T.W., Brodeur, R.D., 2005. Community structure of near-surface zooplankton in the northern California Current in relation to oceanographic conditions. Deep-Sea Research II, this issue [doi:10.1016/ j.dsr2.2004.09.027]. Ressler, P.H., Brodeur, R.D., Peterson, W.T., Pierce, S.D., Vance, P.M., Rstad, A.R., Barth, J.A., 2005. The spatial distribution of euphausiid aggregations in the northern California Current during August 2000. Deep-Sea Research II, this issue [doi:10.1016/j.dsr2.2004.09.032]. Schuyler, Q., Sullivan, B.K., 1997. Light responses and diel migration of the scyphomedusa Chrysaora quinquecirrha in mesocosms. Journal of Plankton Research 19, 14171428. Schwing, F.B., Bograd, S.J., Collins, C.A., Gaxiola-Castro, G., Garc a, J., Goericke, R., Gomez-Valdez, J., Huyer, A., Hyrenbach, K.D., Kosro, P.M., Lavaniegos, B.E., Lynn,