NOAA ERL Special Report

Report on the FOCI International Workshop on

Recent Conditions in the Bering Sea

Seattle, Washington, November 9–10, 1998

S.A. Macklin (editor)

Pacific Marine Environmental Laboratory

7600 Sand Point Way NE

Seattle, WA 98115

January 1999

Contribution 2044 from NOAA/Pacific Marine Environmental Laboratory

Contribution B358 from Fisheries-Oceanography Coordinated Investigations
NOTICE

Mention of a commercial company or product does not constitute an endorsement by

NOAA/ERL. Use of information from this publication concerning proprietary products or the

tests of such products for publicity or advertising purposes is not authorized.

Contribution No. 2044 from NOAA/Pacific Marine Environmental Laboratory

Contribution B358 from Fisheries-Oceanography Coordinated Investigations

For sale by the National Technical Information Service, 5285 Port Royal Road

Springfield, VA 22161

ii
Contents iii

Contents

Executive Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Climate and Upper Ocean Physics . . . . . . . . . . . . . . . . . . 5

Atmosphere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Ocean . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Nutrients and Lower Trophic Level Response . . . . . . . . . . . 13

Lower Trophic Level Studies . . . . . . . . . . . . . . . . . . . . . . . 13

Nutrient Cycling and Phytoplankton Growth . . . . . . . . . . . . . 13

Comparisons of Recent Spring Blooms . . . . . . . . . . . . . . . . . 14

Coccolithophore Bloom . . . . . . . . . . . . . . . . . . . . . . . . . 14

Zooplankton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

Interannual Variability in Zooplankton Abundance . . . . . . . . . . 17

Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

Living Marine Resources (Groundfish, Salmon, and Crabs) . . . 21

Seabirds and Marine Mammals . . . . . . . . . . . . . . . . . . . . 27

Seabirds and Marine Mammals . . . . . . . . . . . . . . . . . . . . . 27

Cetaceans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

Conclusions of Working Groups . . . . . . . . . . . . . . . . . . . . 31

Recommendations . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

Appendix A: Attendees . . . . . . . . . . . . . . . . . . . . . . . . . 38

Appendix B: Agenda . . . . . . . . . . . . . . . . . . . . . . . . . . 44

Appendix C: Discussion Group Assignments . . . . . . . . . . . . 47

List of Figures

1 The Bering Sea. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2 The eastern Bering Sea shelf is larger than California. . . . . . . . . 3

3 Sea level pressure anomalies, in millibars, during early summer 1997

(top) and 1998 (bottom). . . . . . . . . . . . . . . . . . . . . . . . . 6

4 Pacific Ocean sea surface temperature anomalies, in degrees Celsius,

during early summer 1997 (top) and 1998 (bottom). . . . . . . . . . 7

5 Time series of the Pacific Decadal Oscillation (PDO). . . . . . . . . 8

6 Hydrographic regimes of the southeastern Bering Sea shelf. . . . . . 10

7 Near sea surface temperature (top) and depth averaged temperature

(bottom) at mooring site 2. . . . . . . . . . . . . . . . . . . . . . . . 12

8 Spring abundance and stage composition of C. marshallae over the

middle shelf domain. . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

9 Densities of adult euphausiids in and around the inner front region. . 20

10 Recruitment trends of some Bering Sea groundfish from the 1998

groundfish stock assessments. . . . . . . . . . . . . . . . . . . . . . . 22

11 Eastern Bering Sea pollock average individual growth by year as observed from the fishery. . . . . . . . . . . . . . . . . . . . . . . . . . 24

Report on the FOCI International Workshop on

Recent Conditions in the Bering Sea

S.A. Macklin (editor)

Executive Summary

Indications of unusual conditions in the Bering Sea during the summers of

1997 and 1998 included extensive die-offs of seabirds, rare coccolithophore

blooms (a type of non-toxic, microscopic, marine plant), unanticipated low

salmon runs, warm sea surface temperatures, and altered ocean currents

and atmospheric conditions. These changes prompted NOAA FisheriesOceanography Coordinated Investigations (FOCI) scientists to convene an

international workshop with more than 70 participants from a broad range

of disciplines and geographic areas. This ecosystem is of particular concern

because the eastern Bering Sea provides almost half of the fish and shellfish caught in the United States. The implications of these environmental

changes on management of living marine resources was a prominent theme

during the workshop on November 9–10, 1998, in Seattle, Washington.

At the workshop it was established that the springs and summers of 1997

and 1998 were in fact very unusual in a number of ways. The most direct human impact of these conditions was the unanticipated low returns of sockeye

and chinook salmon. Although escapement goals of these fisheries were met,

the low commercial and subsistence catches created considerable economic

hardship. Atmospheric conditions over the eastern Bering Sea in spring

and summer 1997 included light winds, warm temperatures, and clear skies.

These caused a shallow mixed layer; nutrients above the pycnocline were

quickly depleted, leading to the largest coccolithophore bloom ever seen in

the area. The links between these events and the low salmon runs and other

ecosystem anomalies such as the extensive seabird die-offs, remain unclear.

Although spring of 1998 was much stormier than 1997, and stratification and

the spring bloom were delayed, a coccolithophore bloom occurred again, as

did low salmon runs.

It became apparent that the Bering Sea ecosystem is a complex and

highly variable environment. Atmospheric forcing on several temporal scales

(e.g., Pacific Decadal Oscillation, El Ni ̃no, Global Climate Warming) affects

the physical oceanography of the area. In turn, marine forcing causes effects

that cascade through the ecosystem, although we do not understand the

exact mechanisms. The sum of all of these influences produces the conditions

observed at any given time. Our understanding of this system is limited by

lack of historical data.

Recommendations from the workshop included long-term monitoring of

key physical and biological parameters at a number of locations in the eastern Bering Sea. Establishing the linkages between the physics and biology

of the area will require effective interdisciplinary research in a variety of
2 S.A. Macklin (ed.)

Bering Sea

North Pacific Ocean

Gulf of

Alaska

Bristol

Bay

Alaska

A l e u t i a n I s l a n

d s

Bering Strait

Pribilof

Is.

St. Lawrence I.

St. Matthew I.

Alaskan Peninsula

Kodiak

I. Site 2

Russia

Koryak

Anadyr

Bay

Chukotskiy

Peninsula

Kamchatka Strait

C. Olyutorskiy

Kamchatka

Peninsula

Yukon R.

Kuskokwim R.

C. Newenham

Figure 1: The Bering Sea.

modes: modeling, process-oriented, monitoring, and retrospective. Mechanistic understanding of the recruitment processes of a number of key species

is needed. In some cases we have seen correlations between climatic trends

and recruitment trends, but do not know the mechanisms responsible for the

correlations. It is crucial to know the mechanisms because experience has

taught us that correlations break down when the system is altered dramatically.

With our present level of understanding, we will not be able to predict

events such as the unusual occurrences of 1997–1998. This will hamper

effective management of living marine resources in this productive region.

Introduction

There were many indications of unusual conditions in the Bering Sea (Figure 1) during the summers of 1997 and 1998: extensive die-offs of seabirds,

rare coccolithophore blooms, unanticipated low salmon runs, warm sea surface temperatures, and altered ocean currents and atmospheric conditions.

In response to the occurrence of these anomalies, Fisheries-Oceanography

Coordinated Investigations (FOCI) directors Drs. Arthur Kendall, Jr.

(NOAA Alaska Fisheries Science Center [AFSC]) and Phyllis Stabeno

(NOAA Pacific Marine Environmental Laboratory [PMEL]) convened a 2-

day international workshop at the Sand Point NOAA facility, Seattle, Washington, on November 9 and 10, 1998.

The purpose of the workshop was to share information, integrate knowledge, suggest mechanisms, propose hypotheses, and outline future research
FOCI International Workshop Report 3

Figure 2: The eastern Bering Sea shelf is larger than California.

needs to address and understand changing conditions in the Bering Sea. This

ecosystem is of particular concern because the eastern Bering Sea provides

almost half of the fish and shellfish caught in the United States. Most of the

catch comes from the continental shelf that borders Alaska’s western coast,

an area much larger than the state of California (Figure 2). Besides producing abundant fish and shellfish, this region also supports large numbers of

resident and migratory seabirds and marine mammals. A prominent theme

during the workshop was the implications of recent environmental changes

on the management of these living marine resources.

Several thematic questions served as a focus for participants at the workshop, particularly the keynote speakers: In your field of expertise, how were

conditions in the eastern Bering Sea during 1997 and 1998 different from

usual? What mechanisms caused these anomalies? Is there evidence that

unusual conditions will persist? What are the implications for the future of

the ecosystem and its living marine resources?

The organizing committee for this workshop comprised Mr. Allen Macklin (PMEL): atmosphere and ocean physics; Dr. Jeffrey Napp (AFSC): lower

trophic levels; Dr. Cynthia Tynan (Joint Institute for the Study of Atmosphere and Ocean [JISAO]): upper trophic levels, except fish; Dr. Richard

Brodeur (AFSC): fish and fisheries.

Participants at the workshop included a member of the fishing indus-
4 S.A. Macklin (ed.)

try, representatives of fisheries and scientific management, various scientific

disciplines, environmental organizations, and native Alaskan groups (Appendix A). From the agenda (Appendix B), the Monday session featured

eight 20-minute keynote presentations detailing physical and biological observations made during the last 2 years and identifying linkages to other

parts of the Bering Sea system and to global weather/oceanographic patterns. Following the keynote addresses, other attendees, including those

from Canada, Japan, and Russia, had the opportunity to present findings.

A reception featuring an address by a member of the fishing industry, Mr.

Bart Eaton, Vice President of Trident Seafoods Corporation, was held Monday evening. Mr. Eaton stated that industry, as well as management, needs

to be flexible in responding to the changing conditions of the resources in

the Bering Sea. He also challenged scientists to find new ways for industry

to assist with data collection.

On Tuesday morning, participants broke into two groups to discuss and

answer the following questions:

• What are the physical-chemical-biological mechanisms causing the

changes in the ecosystem observed in 1997 and 1998?

• What research is needed to better understand the origin and persistence of these changes?

• Given the large changes in the Bering Sea ecosystem, how do we manage marine resources in a fluctuating and highly variable environment?

Products from the workshop included a NOAA press release (12/30/98),

this summary report with recommendations and endorsement of the draft

Science Plan for the Bering Sea Ecosystem, and an Internet-based chat room

to facilitate further discussion on the issues covered by the workshop.
FOCI International Workshop Report 5

Climate and Upper Ocean Physics

Contributors: P.J. Stabeno, N. Mantua, J.E. Overland, S.A. Macklin, and

G. Weller

Atmosphere

The Bering Sea (Figure 1) is highly responsive to meteorological and climatological forcing. Local radiative warming and cooling, thermal advection,

and wind mixing occur through processes associated with the progression of

extra-tropical storms through the region. Storm tracks, frequencies, and intensities for the Bering Sea are affected by teleconnections to equatorial (e.g.,

El Ni ̃no-Southern Oscillation [ENSO]) and other climate forcing. Additionally, decadal-scale climate patterns, as demonstrated by various atmospheric

indices (e.g., Pacific North American Index [PNA], Pacific Decadal Oscillation [PDO]) provide quasi-stable weather regimes. Finally, global warming

may be a long-term climate trend affecting the Bering Sea region.

In 1997, the early summer (May–June–July [MJJ]) climate over the

North Pacific Ocean and Bering Sea was uncharacteristically warm, windless, and cloud free. These benign weather conditions followed a relatively

mild winter with anomalously low snow pack in much of western Alaska. A

persistent pattern of anomalously high sea level pressure (SLP) (shown in

the upper panel of Figure 3) over the Aleutian Islands and Bering Sea, with

regions of anomalously low SLP to both the southwest and southeast of the

Aleutians, contributed to these meteorological events. Such regions of high

SLP anomalies are typically associated with anomalously fair weather, while

low SLP cells are typically associated with anomalously stormy weather.

In contrast, meteorological conditions observed in the late spring–early

summer of 1998 were characterized by frequent storms, strong winds, and

relatively cloudy skies. The SLP anomaly patterns for MJJ 1998 (lower

panel of Figure 3) were nearly opposite of those observed in the same calendar months of 1997. It is also interesting to note that in each of the past

two summers, anomalous SLP and circulation patterns have been relatively

strong (though very different) across the North Pacific. The anomalous

weather during the past two spring and summer seasons has contributed to

relatively large sea surface temperature (SST) anomalies as well. In MJJ

1997 extreme El Ni ̃no conditions were well underway in the tropics. The

characteristic warming centered on the equator in the eastern and central

equatorial Pacific was a dominating feature of Pacific SST anomalies (upper

panel of Figure 4). Concurrent with equatorial warming, SSTs in the Northeast Pacific, Gulf of Alaska, and Bering Sea warmed at equally impressive

rates. By MJJ 1998 the pattern of Pacific SST anomalies had changed dramatically (lower panel of Figure 4). The first signs of the transition from El

Ni ̃no to La Ni ̃na conditions were evident as temperatures cooled sharply in

the central equatorial Pacific. Likewise, North Pacific SSTs were markedly

cooler in MJJ 1998 than they were in the same calendar months of 1997,

with only small regions of slightly warmer than average temperatures in the

eastern Bering Sea and coastal waters of the northeastern Pacific. Another
6 S.A. Macklin (ed.)

85N

80N

75N

70N

65N

60N

55N

50N

45N

40N

35N

30N

25N

20N

85N

80N

75N

70N

65N

60N

55N

50N

45N

40N

35N

30N

25N

20N

120E 140E 160E 180 160W 140W 120W 100W 80W

120E 140E 160E 180 160W 140W 120W 100W 80W

Figure 3: Sea level pressure anomalies, in millibars, during early summer 1997 (top) and 1998 (bottom).
FOCI International Workshop Report 7

Figure 4: Pacific Ocean sea surface temperature anomalies, in degrees Celsius, during early summer

1997 (top) and 1998 (bottom).
8 S.A. Macklin (ed.)

1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000

Nov-Mar PDO Index

Figure 5: Time series of the Pacific Decadal Oscillation (PDO).

major change in upper ocean temperatures took place in the central and

western North Pacific. Whereas SSTs were cooler than average in a wide

band off the Asian coast in 1997, by MJJ 1998, SSTs had become 1 to a few

degrees centigrade warmer than average in this same region.

What does the future hold? In the past few months, the trends toward

anomalously cool SSTs in the NE Pacific and warm SSTs in the central

North Pacific have continued. For most of the post-1977 period, PDO index

values have been positive (Figure 5), indicating strong persistence of the

“warm” (positive PDO) regime that appears to have favored such ecosystem

features as high Alaska salmon production and low production of major

crab and shrimp populations in the Bering Sea. From this index, there

is no suggestion of a “regime shift” in the period since 1977. Given the

strong likelihood that La Ni ̃na conditions will persist through the winter

months of 1998–1999, it is also very likely that La Ni ̃na influences on North

Pacific winter climate will favor the maintenance and possible amplification

of negative PDO SST anomalies through (at least) the spring of 1999.

Beyond the spring of 1999, predicting North Pacific climate (both in

the ocean and atmosphere) becomes much more problematic. The present

day understanding of the PDO phenomenon is quite poor, especially when

compared with the state of knowledge regarding ENSO. The synergy between observations, theory and modeling provided for rapid advances in

understanding and predicting ENSO. Compared with our ability to monitor, understand, and predict ENSO variability, PDO science lags many years

behind. Using the ENSO experience of the past two decades as a model, it

seems of utmost importance to develop a comprehensive climate monitoring

system for the North Pacific Ocean. The TOGA/TAO array in the equatorial Pacific has provided much valuable information to ENSO studies and

predictive capability.

Long-term climate trends must also be taken into account in assessing

impacts in addition to interdecadal and ENSO-related climate variability.

From paleoclimatic evidence, it appears that present conditions are highly

anomalous, and it seems very likely that we have already begun to see the

greenhouse effect in the Arctic. Air temperature data show cooling in winter
FOCI International Workshop Report 9

in the western Bering Sea, while the eastern Bering Sea and Alaska show

warming.

Ocean

The Bering Sea forms a buffer between the North Pacific and Arctic Oceans

(Schumacher and Stabeno, 1998; Stabeno et al., in press). It is also a region

of extremes. Physically, it is divided into almost equal parts: a basin (maximum depth ∼4000 m) and a broad (>600 km), shallow shelf on the eastern

side (Figure 1). A defining characteristic of the Bering Sea is the presence

of seasonal sea ice. From June through October the Bering is largely icefree. Typically in November, sea ice begins forming in the polynyas and is

advected southward by arctic winds. In extreme years most of the broad,

eastern shelf is covered in ice; in weak years the sea ice does not extend south

of 60◦N. The melting ice freshens the water and lowers its temperature, contributing markedly to water properties over the shelf. Ice extent has changed

markedly over the last 30 years. The ice extent was typically extensive from

the early 1970s until 1976, reaching a peak in 1976, when it covered much of

the eastern shelf for 2 months. After the regime shift in 1976, the extent and

duration of ice over the shelf diminished. The most extensive ice coverage in

recent years occurred in 1995, when ice covered much of the shelf for about

a month. The ice extent in 1997 was also above average, arriving over the

southern part of the shelf in March and retreating in early April. In contrast,

the ice in 1998 arrived during January and retreated during February.

During winter, the water over the southeast shelf is usually well mixed

to a depth of ∼100 m, while during summer, three distinctive hydrographic

regimes are evident (Figure 6), as described by the Processes and Resources

of the Bering Sea Shelf (PROBES) project. The “coastal domain” (water

depth <50 m) is well mixed or weakly stratified from a combination of tidal

and wind mixing. The “middle shelf domain” (50 m < water depth < 100 m)

is two layered. An upper, wind-mixed layer is separated from a tidally

mixed bottom layer by a strong pycnocline. The “outer shelf domain” is

characterized by a wind-mixed surface layer and tidally mixed bottom layer

separated by a region of finestructure with gradually increasing density. The

coastal domain and the middle shelf domain are separated by the “inner

front,” and the middle and outer shelf domains are separated by the “middle

transition zone.” Support from the National Science Foundation’s Office of

Polar Programs Inner Front project permitted a careful investigation of the

dynamics and structure of the inner front during 1997 and 1998. During

1997, weak winds resulted in stratification extending the middle shelf domain

coastward to a depth of ∼30 m. In contrast, the strong winds of 1998 resulted

in the coastal domain extending out to ∼70 m until June, at which time the

winds weakened and the more typical hydrographic regimes appeared.

A series of FOCI biophysical moorings have been deployed at a location

near the center of the middle shelf domain (Site 2; Figure 1) for the last

4 years. These moorings were maintained with support from two NOAA

Coastal Ocean Program (COP) regional projects: Bering Sea FOCI and

Southeast Bering Sea Carrying Capacity (SEBSCC). This is the only location
10 S.A. Macklin (ed.)

9

Salinity (psu)

Temperature (°C)

0

50

100

150

0

50

100

150

0

50

100

25.0 26.0 24.0 25.0 26.0

Tidally and

wind stirred

~homogenous

24.0 25.0

Z< 50 m

100 < Z< 50 m Tidally stirred

Wind stirred Fine structure

Vertical Structure

Sigma T 100 < Z < 180

33.2

33.0

32.8

32.6

32.4

32.0

32.2

31.8 31.6

31.4 31.2

30.6

4

6

7

9 10

7

4

5

<2 2

30.4

Slope water

Front

600 km 500 km 400 km 300 km 200 km 100 km 0 km

8

Middle Shelf

Middle transition zone Inner Front

Coastal

5

8

Outer shelf

Shelfbreak

Figure 6: Hydrographic regimes of the southeastern Bering Sea shelf.

where long-term water column measurements have been made in the Bering

Sea. In this discussion we will focus on this location, since it is ideal for

examining direct warming of the water column in the spring and summer.

The weak winds and cloud-free days of 1997 resulted in a shallow mixed layer

with warm SST. The temperature (measured at 1–2 m below the surface;

upper panel of Figure 7) shows a steady warming during each year, beginning

in April with maximum temperatures occurring in late July or early August.

The x’s are historical temperature observations near this location dating

back to 1962. The warmest temperatures ever recorded over the middle

shelf occurred in 1997.

The wind forcing was at the opposite extreme in 1998 compared to 1997.

In 1998, strong winds resulted in a well-mixed water column well into May.

The water column did not stratify until June. In July the winds weakened,

resulting in a shallow mixed layer (15 m) and warm near surface temperatures. The warm upper water temperatures in 1998 were thus a result of

warmer than usual temperatures in April (the initial condition of the heating cycle) and shallow mixed layer in mid July. The warm temperatures
FOCI International Workshop Report 11

in April resulted from the atmospheric conditions in conjunction with ice

cover. When ice was present over the southeastern shelf at site 2, the winds

were weak, thus the cooling resulting from ice melt was limited to the upper

30 m. A reservoir of relatively warm water remained at depth. The storms

of March and April stirred the water column, and the near-surface ocean

temperature increased due to the mixing of warmer, deeper water. This is

seen in the average vertical temperature at site 2 (lower panel of Figure 7).

The average temperature (and thus heat content) of the water column in

1998 was higher than in 1997, and was similar to that observed in the warm

years of 1978, 1979, and 1981. The implications of this for the coming year

are not known, but it is interesting to note that in late November 1998, the

Bering Sea is ice free and SSTs are above 6◦C over part of the southeastern Bering Sea shelf. Thus the system enters the 1998/1999 winter season

with higher than average heat content. To remove the heat and bring the

system back toward its state in the early part of this decade will require

above-average sea ice production and extended duration of arctic winds.

In addition to extremes that have occurred on the southeastern shelf,

changes of transport through Bering Strait have occurred. Over the last

several years, there appears to have been both a reduction in the total northward transport from the Bering Sea into the Arctic and in its salinity. Both

changes could have repercussions on the Arctic Ocean, since Bering Strait is

the only connection between the Pacific Ocean and the Arctic. The decrease

in transport could be a result of changing wind directions or a decrease in

the difference in the height between the Pacific and Arctic Ocean. The decrease in salinity resulted either from a decrease in the amount of sea ice

(and hence a decrease in the amount of brine rejection) or a decrease in the

salinity of the water on the shelf. There is some indication that over the last

few years there has been a decrease in salinity over the southeastern shelf,

but the variability of this signal is large.

Conclusions

Because observations have been limited over the years, it is difficult to determine whether recent extremes in the physical parameters of the Bering Sea

are within the bounds of historical variability, the result of a decadal-scale

regime shift, or due to global climate change. Much of our insight has been

a result of the ∼5-yr time series obtained from a single location over the

Bering Sea shelf.

• Anomalously weak winds occurred over the Bering Sea in 1997, and

unusually strong storms occurred during 1998.

• Above normal SSTs were observed during the summer of 1997 and

1998, with the warmest temperatures occurring in 1997, but the highest

water column heat content during 1998.
12 S.A. Macklin (ed.)

Figure 7: Near sea surface temperature (top) and depth averaged temperature (bottom) at mooring site 2.
FOCI International Workshop Report 13

Nutrients and Lower Trophic Level Response

Contributors: J.M. Napp, K.O. Coyle, T.E. Whitledge, D.E. Varela, M.V.

Flint, N. Shiga, D.M. Schell, and S.M. Henrichs

Lower Trophic Level Studies

The eastern Bering Sea shelf is very large, larger than the state of California

(Figure 2). To put past and present sampling into perspective, imagine a

scientist using a butterfly net or empty soda bottle to sample water and

animals from the state of California. All sampling is conducted at night or

during periods of thick fog so that the scientist cannot see what is being

sampled. In addition, the research “platform” is a tricycle, so the area

sampled is relatively small. This example is analogous to scientists sampling

the Bering Sea shelf from slow moving ships where the target organisms lie

below the surface of the ocean and cannot be seen. Thus the community must

recognize that many of our generalizations about plankton refer to specific

areas and that a comprehensive examination of pattern and processes is

lacking.

Nutrient Cycling and Phytoplankton Growth

During the late 1970s and early 1980s, PROBES provided a sketch of nutrient cycling, variability, and phytoplankton growth for the eastern Bering

Sea (Whitledge et al., 1986; Whitledge and Walsh, 1986). The annual nutrient cycle was carefully delineated over 4–5 years with regard to vertical

structure along a single cross-shelf transect. Major events in the nutrient

cycle are nutrient replenishment through advection (lateral) or wind-driven

mixing (vertical), and utilization by phytoplankton. Careful observation of

the spring phytoplankton bloom showed that the bloom started offshore and

slowly progressed inshore, across the shelf. The peak in chlorophyll concentration started in mid-April offshore and by early June was observed

over the coastal domain. Changes in the stability of the water column

(Sigma-t) best explained this pattern. The annual cycle of nitrate (NO3)

follows phytoplankton dynamics. Nitrate was replenished over the shelf in

late winter (March) by advection from the basin. Rapid decline of nitrate

in the mixed layer was observed during the spring phytoplankton bloom.

During the summer, phytoplankton growth was driven by regenerated nitrogen (ammonia [NH4]) except when episodic wind mixing events replenished

nitrate in surface waters. These events were shown to introduce 15–20 μM

m−2 of new nitrate into the euphotic zone. This is probably the key to

maintaining a summer population of diatoms.

During 1997 and 1998, different patterns of nutrient utilization with

depth were observed. In 1997, stratification began early due to fewer storms

than usual; 1998 was exactly the opposite, spring was extremely windy and

the onset of stratification was delayed. Normally, uptake of nitrate by phytoplankton shows relatively large values at the surface, decreasing with depth.

In 1997, however, the data clearly indicate that nitrate uptake increased
14 S.A. Macklin (ed.)

with depth and was maximal below the pycnocline. This resulted in depletion of nitrate (and silicate) from the water below the pycnocline (a nutrient

reservoir). Thus more of the nitrate pool was available to phytoplankton

production. In 1998 high nitrate uptake values at the surface were observed,

but the wind mixing was so strong that light limitation prevented a large

spring bloom (accumulation of plant biomass). The result was a slow, rather

than rapid, decrease in nutrient concentration beginning in April through

May and part of June. The consequence was the same as 1997: more of the

nitrate pool was available to the phytoplankton. Preliminary calculations

indicate that primary productivity calculated from nitrate disappearance

during PROBES was 10–30% smaller than calculated for 1997 and 1998.

The end result on system carrying capacity, trophic transfer, or the amount

of the production retained in the water column is not known.

Comparisons of Recent Spring Blooms

The spring phytoplankton bloom is the dominant production event over the

southeastern Bering Sea shelf. At FOCI’s middle shelf domain biophysical mooring, continuous measurements of chlorophyll absorbance have been

recorded (February–September) from 1995 to the present as part of a collaboration between NOAA and academic scientists (Oregon State University, 1995–96; Dalhousie University, 1997–98). These measurements provide

insight into how interannual variability in weather affects the annual production cycle over the shelf (J. Cullen, personal communication). Briefly,

1995 was a heavy ice year, the bloom began early under the ice, and a large

amount of chlorophyll accumulated in the surface waters before presumably

settling to the bottom (Stabeno et al., 1998). Surface nutrients were exhausted at the end of the bloom. In 1997 we observed a similar pattern,

except the ice retreated rapidly and the maximum amount of chlorophyll

was lower than in 1995. Sea ice did not extend onto the southeast shelf

in 1996. Water column stratification and the spring phytoplankton bloom

began much later in 1996 than either 1995 or 1997. In 1998 accumulation of

chlorophyll also occurred very late (late May, early June). We interpret this

to mean that phytoplankton growth and zooplankton grazing over the middle shelf domain were balanced for most of the spring, potentially resulting

in less flux of phytoplankton to the benthic community.

Coccolithophore Bloom

During summer 1997, waters over large portions of the eastern Bering Sea

shelf were turned chalky aquamarine by a bloom of coccolithophores (Emiliania huxleyi; Vance et al., 1998; Sukhanova and Flint, 1998). This was

the first confirmed report of a large-scale coccolithophore bloom in the eastern Bering Sea, although reflectance algorithms used with NASA’s Coastal

Zone Color Scanner (an ocean color remote sensing tool) suggest that smaller

blooms may have occurred between 1978 and 1984 (Brown and Yoder, 1994).

Phytoplankton species counts around the Pribilof Islands in the early 1990s
FOCI International Workshop Report 15

revealed low, non-bloom concentrations of E. huxleyi (M. Flint, personal

communication).

Coccolithophores are small, flagellated photosynthetic cells (approx.

10 μm diameter) covered with calcareous plates that scatter light producing

the aquamarine color. High reflectance and underwater light extinction are

associated with coccolithophore blooms. By September of 1997, the bloom

covered 2.1 × 105 km2 of the eastern Bering Sea shelf (Vance et al., 1998).

Coccolithophore cell densities around the Pribilof Islands in October were

approximately 2.1 × 106 liter−1 and coccolithophores constituted 99.2% of

the phytoplankton biomass (M. Flint, personal communication). Diatoms,

not coccolithophores, dominate the normal fall community. The coccolithophore bloom returned in the early spring of 1998 and remained through

the summer.

What caused the bloom and allowed coccolithophores to replace diatoms

as the dominant summer and fall species? Nutrient uptake experiments

conducted in 1997 indicated that diatom uptake responded much more to

nitrate concentration than did uptake by communities dominated by coccolithophores (J. Goering, personal communication). The f-ratio (nitrate

uptake/uptake of all dissolved inorganic nitrogen species) was 0.2–0.3 for

the diatoms and about 0.1 for coccolithophores. During 1997 and 1998 ammonium concentrations were high on the shelf, nitrate and silicate were low,

and the water temperature was anomalously warm. These conditions probably allowed the coccolithophores to outcompete the diatoms in 1997. We

do not yet know what happened in 1998; ambient nutrient concentrations

and nutrient uptake experiments are being analyzed.

Species of phytoplankton respond differently to the availability of nutrients and trace metals in the water column. In order to understand why

extensive coccolithophore blooms have developed in the Bering Sea, it is important to identify which factors promote their growth. The coccolithophore

E. huxleyi, isolated from Ocean Station “P” (OSP) in the subarctic Northeast Pacific Ocean, appears to grow equally well on different nitrogen sources,

either ammonia or nitrate, in laboratory cultures (D. Varela, personal communication). However, when ammonium is added to nitrate-grown cells, the

uptake rate of nitrate is dramatically inhibited and growth continues. Such

behavior indicates that this coccolithophore may be growing on ammonium

due to the constant availability of this nitrogen source from zooplankton excretion and remineralization. E. huxleyi can also grow in either high or low

iron concentrations and adapts to low iron concentrations by reducing cell

size. Therefore, there is no difference in sinking rates between cells in ironreplete or iron-stressed conditions because under lower iron concentrations

the cells are smaller and sink more slowly. In contrast, the diatom Actinocyclus sp., also isolated from OSP, increases its sinking rate when under iron

stress. In the laboratory, ammonia-grown diatom cells showed signs of energetic advantage over nitrate-grown cells when under iron stress. Thus, under

the iron-limited scenario of the subarctic Northeast Pacific, diatoms may be

growing on ammonium.

These laboratory results are in good agreement with field observations in

the subarctic Northeast Pacific. Studies along a transect extending from the
16 S.A. Macklin (ed.)

continental slope off the southwestern corner of Vancouver Island (British

Columbia, Canada) to OSP showed that most of the primary production is

based on ammonia, and is due to small cells (<5 μm) well adapted to the

low ambient iron levels. It is possible that nitrogen and iron limitation on

the southeastern Bering Sea shelf would favor the growth of E. huxleyi over

diatom species. At present there is no data on iron availability or limitation

on the Bering Sea shelf.

Zooplankton

Zooplankton is an important link in the food web between primary production and apex predators (fish, seabirds, and marine mammals). There

are many different types and forms of zooplankton. The key, when looking

for system anomalies, is to examine types or species that might be sensitive, either through physiology (e.g., temperature) or life history, to the

observed system anomalies. Many zooplankton taxa that are prey to fish,

birds, and mammals (e.g., large copepods and euphausiids) have annual life

cycles. Thus biological or environmental influences experienced by a population during one season may be observed both during that season, during

subsequent seasons, and during the following year. For example, high predation on female zooplankters or severe conditions during spring may mean

less reproduction and fewer juvenile zooplankton available during the summer as prey. Poor feeding conditions during the summer may impact those

invertebrates that rely on lipid stores to make it through the long winter,

thus decreasing the brood stock for the following year.

Each of the three distinct hydrographic regimes has its own zooplankton fauna (Cooney and Coyle, 1982; Vidal and Smith, 1986; Smith, 1991).

The outer shelf domain is dominated by large oceanic copepods (Neocalanus

spp.) and one euphausiid (Thysanoessa inermis). The zooplankton biomass

of the middle shelf domain is dominated by a moderately sized copepod

(Calanus marshallae) and another euphausiid (T. raschii), and the coastal

domain is dominated by small copepods (Acartia spp. and Oithona spp.).

Secondary production among the three regimes is highest in the outer shelf

domain and lowest over the coastal domain. Primary production and grazing are usually balanced in the outer shelf domain, but are largely unequal

in the middle shelf and coastal domains. Often the phytoplankton blooms

sink to the bottom in the latter two domains, providing energy for a rich

benthic community. Between each domain lies a front or transition zone.

Secondary production at the shelf edge, another transition zone, is thought

to be greater than that of the three domains due to persistent upwelling of

nutrients to the euphotic zone (Springer et al., 1996). Production at the

inner front is currently being studied by the Inner Front project (G. Hunt,

personal communication). Nearly continuous supply of nutrients plus active

concentration of prey at these fronts appear to make the areas loci for apex

predators such as seabirds and marine mammals.

Long-term (decadal and longer period) fluctuations in zooplankton biomass of the eastern Bering Sea have been noted. Zooplankton biomass was

highest in the 1960s (800–900 mg m−3) and decreased in the 1970s to 1980s
FOCI International Workshop Report 17

and again in the early 1990s to 200–300 mg m−3 (Sugimoto and Tadokoro,

1997). The same authors studied the relationship between long-term anomalies in summer zooplankton biomass and winter winds, summer phytoplankton, and Asian pink salmon catches. They concluded that predation pressure by salmon controls the year-to-year variability in summer zooplankton

biomass, but bottom up control (winds to phytoplankton to zooplankton)

was more influential than top down control on longer (decadal) time scales.

A long-term increase in jellyfish biomass has also been noted for the eastern Bering Sea. Since 1990, jellyfish (Scyphomedusae) biomass in standard

trawl surveys has tripled (Brodeur et al., submitted). Jellyfish are carnivorous and increased biomass may mean increased predation on juvenile fish

and increased competition with juvenile fish for zooplankton prey.

Interannual Variability in Zooplankton Abundance

First through Bering Sea FOCI and presently through SEBSCC, FOCI has

been collecting samples for zooplankton biomass and community composition in larval and juvenile pollock habitat on the southeastern Bering Sea

shelf. Collections are made in the winter, spring, and fall. Recent springtime abundance and stage composition of C. marshallae over the middle

shelf domain show a pattern that may reflect interannual variability in water temperature and timing of the spring bloom (Figure 8). Note that C.

marshallae nauplii are consumed by larval fish, and copepodites are important dietary components of age-0 fish. The highest abundances and earliest spawning (deduced from the early appearance of young developmental

stages) was during 1995, the year with the coldest water temperatures and

the earliest spring bloom (J. Napp and C. Baier, personal communication).

Springtime abundances in 1996 and 1997 were comparable, but the appearance of young developmental stages was latest in the warm year (1996) when

the spring bloom was delayed. Analysis of FOCI 1998 samples (also a warm,

late bloom year) is ongoing. Japanese researchers report that summer zooplankton biomass in 1998 was neither anomalously high nor low (N. Shiga,

personal communication). The extremes in recent years are not directly

comparable to a published comparison of zooplankton from PROBES: 1981

was warm and calm, while 1980 was cool, but stormy (Smith and Vidal,

1986).

Following cohorts of C. marshallae to the end of summer, FOCI has found

that if the year starts with relatively more females and copepodites, then

relatively more are found in that year class at the end (J. Napp, personal

communication). When the years were ranked by abundance of C. marshallae sampled around the Pribilof Islands during September, the rank order

of the years was 1995, 1997, and 1996 (highest to lowest). The exception

to this was the 1996 year class. Both 1996 and 1997 began the years with

approximately the same abundance of C. marshallae, although reproduction

began much later in 1996. In 1996, however, relatively high abundances of

age-0 pollock were found over the southeastern shelf (Wilson and Brodeur,

1998), so low abundances of C. marshallae may also have been due to predation by age-0 pollock (N. Shiga, personal communication). Thus at this
18 S.A. Macklin (ed.)

Figure 8: Spring abundance and stage composition of C. marshallae over the middle shelf domain.
FOCI International Workshop Report 19

Cape Newenham

Dillingham

Nunivak

Port Moller

Slime Bank

Abundance

(No. m-2)

Jun-97

Sep-97

Jun-98

0

50

100

150

200

250

300

P < 0.001

P < 0.05

Figure 9: Densities of adult euphausiids in and around the inner front region.

point in our analyses, we do not see either a sharp increase or decrease in

abundance of one of the dominant zooplankton species of the middle shelf

domain, the one most affected by the anomalously high water temperatures

and coccolithophore bloom.

The most notable distinction between 1997 and other years in zooplankton abundance and distribution were the significantly lower densities of adult

euphausiids in and around the inner front region (Inner Front project, K.

Coyle, personal communication; Figure 9). Short-tailed shearwaters, a migratory sea bird that relies on surface swarms of euphausiids for food, suffered high mortalities during the summer of 1997 (G. Hunt, personal communication). Those shearwaters surviving to the end of summer had broadened

their diet to include squid and juvenile fish. Interestingly, adult plus juvenile

euphausiid abundance measured around the Pribilofs (including the inner

front) in 1997 by SEBSCC (Napp, unpublished; Brodeur, unpublished) was

higher than that measured in either 1995 or 1996. Investigators from the

Inner Front project and SEBSCC are currently comparing their zooplankton

data sets for anomalies in biomass, distribution, and community composition

that may be attributed to recent environmental anomalies.
20 S.A. Macklin (ed.)

Conclusions

• Interdecadal variations in zooplankton biomass in the southeastern

Bering Sea are controlled by physics and phytoplankton concentration.

• Sea ice and wind mixing affect the timing and magnitude of the spring

phytoplankton bloom.

• The abundance of selected zooplankton taxa appeared to respond to

variations in the timing and magnitude of the spring bloom.

• Interannual differences in the distribution of zooplankton (as prey for

apex predators) are important.

• In the fall of 1997, euphausiid prey for diving seabirds existed around

the Pribilof Islands, but not in the coastal domain or inner front adjacent to the mainland. This resulted in massive mortalities for shorttailed shearwaters.

• In the summer of 1998, age-0 pollock and oceanic zooplankton were

found at the inner front, suggesting large-scale, shoreward transport

of animals from at least the outer shelf domain.
FOCI International Workshop Report 21

Living Marine Resources (Groundfish, Salmon,

and Crabs)

Contributors: R.D. Brodeur, G.H. Kruse, P.A. Livingston, G. Walters, J.

Ianelli, G.L. Swartzman, M. Stepanenko, and T. Wyllie-Echeverria

The most visible sign of unusual fisheries events in the Bering Sea was the

surprisingly low return of sockeye salmon to Bristol Bay in 1997 and 1998.

This is the world’s largest wild sockeye fishery. Actual catch was 49% and

47% of that forecast in 1997 and 1998, respectively. Although salmon escapement goals were met for this stock, there were substantial economic

hardships incurred and subsistence harvest needs were not met. The State

of Alaska has taken emergency measures to ease the hardships. In addition, salmon runs to the Yukon River, Kuskokwim River, and several other

river systems, were well below predicted levels (Kruse, 1998). The Yukon

River chum run was weak in 1998, and the commercial catch (about 4% of

the mean catch since 1974) was restricted to bycatch in the early chinook

salmon openings. Anecdotal evidence suggests the fish that did return to

the Yukon River were smaller than normal, which is somewhat unusual for

a small run of salmon. Larger than average fish are usually associated with

low returns. The decrease in abundance and size was noted across several

groups, which suggests that the causative factor was likely to be of marine

origin (Kruse, 1998). Despite these shortfalls in the Bering Sea, returns of

pink salmon in the Gulf of Alaska were generally strong. The statewide catch

of pink salmon (104 million) was the fifth highest ever. Taken together, the

total commercial catch of Alaska salmon was 150 million fish, which was the

eighth highest ever recorded. The mixed returns may indicate that some

regional factor associated with the Bering Sea, such as increased temperatures, more predation, or reduced food availability, may be affecting these

fish. Marine factors affecting survival are generally thought to be the most

important during early marine life, whereas later, marine factors can have

significant impacts on growth. However, at this point in time, we are unable

to disentangle the contributions of early marine conditions from the unusual

ocean conditions of the last 2 years on reduced salmon runs in 1997 and

1998. So, we cannot yet tell if the impacts of the last 2 years will carry over

into future return years, even if ocean conditions return to normal in 1999.

Groundfish stocks in the eastern Bering Sea are surveyed each summer by

the AFSC, and stock assessments are provided annually to the management

agencies. From results of the most recent (1998) assessments, there do not

appear to be any anomalies observed in the distribution patterns of the dominant fish species that could be associated with the unusual ocean conditions

in 1997 and 1998. Long-term trends in biomass and recruitment for some

species have been assessed. For example, yellowfin sole, rock sole, flathead

sole, and arrowtooth flounder show a recent increase in adult biomass due

mainly to good year classes in the late 1980s and early 1990s. In contrast,

other species, such as Pacific cod, Alaska plaice, and Greenland turbot have

had few or no recent strong year classes and have been declining in biomass
22 S.A. Macklin (ed.)

Flatfish recruitment

-

100,000

200,000

300,000

400,000

500,000

600,000

700,000

800,000

900,000

6971737577798183858789919395

Year class

Arrowtooth flounder (age 2

number -1000s)

-

5,000

10,000

15,000

20,000

25,000

30,000

35,000

40,000

Greenland turbot

(age 1 number -1000s)

Arrow tooth flounder

Greenland turbot

Gadid recruitment

0

200

400

600

800

1000

1200

1400

19781980198219841986198819901992199419961998

Year

Age 1 cod recruits

(millions)

0

10,000

20,000

30,000

40,000

50,000

60,000

70,000

Age 1 pollock recruits

(billions)

Pacific cod

Walleye pollock

Figure 10: Recruitment trends of some Bering Sea groundfish (arrowtooth flounder, Greenland turbot,

walleye pollock, and Pacific cod) from the 1998 groundfish stock assessments.
FOCI International Workshop Report 23

throughout the 1990s. Walleye pollock show strong pulses of recruitment and

the biomass has fluctuated over the last 35 years. The 1996 year class appears

to show the most recent above-average recruitment. This strong year class

was detected in age-0 surveys of walleye pollock conducted in mid-summer

in cooperative surveys with Japanese scientists (Wilson and Brodeur, 1998).

An interesting observation is that the recruitment of some flatfish species

vary out of phase with each other (e.g., Greenland turbot and arrowtooth

flounder, yellowfin sole and rock sole), whereas recruitment of gadid stocks

(walleye pollock and Pacific cod) vary roughly in phase (Figure 10). The various groundfish have been classified into trophic guilds based on their dietary

preferences. The inshore benthic infauna consumer guild (soles and plaice)

have shown a general increase since 1979, while the fish and crab consumer

guild (sculpins, cod, skates, and halibut) have been declining since 1988.

Trends in forage fish (herring, capelin, eulachon, juvenile cod, and juvenile walleye pollock) abundance also have been documented by AFSC summer surveys. Most of these forage fishes have declined over the last 5 years.

Although the catch rates in 1997 were not particularly low, 1998 had fairly

low catches for all species but pollock. Both surface and bottom temperatures measured during the survey showed that 1994 and 1995 were relatively

cold years, while 1996 through 1998 were warmer than average. The cold

pool was very extensive in 1995, extending well into Bristol Bay.

Fishery data collected by NMFS observers are very extensive. Length,

weight, and age are determined for a large number of pollock each year.

These data are routinely used in stock assessments for Bering Sea pollock

and can be used to evaluate how average growth in weight changes over time

and age. Results indicate that the annual average weights-at-age from the

fishery are highly variable (Figure 11). Based on analyses of growth by cohort from 1996–1997 (e.g., average weight increase of 4 year-old pollock in

1996 compared to 5 year-olds in 1997), pollock growth during this year was

considerably lower than average. This may reflect changes in the distribution of the fishery, differences in optimum temperature regimes for pollock

growth, or possibly shifts in prey availability and consumption. Analyses

are ongoing to see if lower than average growth persisted through 1998.

In addition to interannual variability, there is a consistent pattern of

weight increase with age. The average annual growth decreases from ages

4 to 6, and then appears to increase again with older ages. Preliminary

examinations of bottom-trawl survey estimates of average weights-at-age

suggest a similar pattern is also present from those samples. This would

indicate that the pattern is not due to the target age-classes of the fishery.

In recent years, Russian research surveys have been conducted throughout the Bering Sea to assess the abundance and distribution of various age

classes of pollock and other fish. During the warm summer and autumn

of 1996, a significant part of the mature eastern Bering Sea pollock population, including the very abundant 1992 year class, inhabited the area to

the northwest of the Pribilof Islands. In the warm spring of 1997, most of

the western Bering Sea pollock spawned on the northwestern Russian shelf.

Juvenile pollock were extremely abundant (5.3 × 109 fish) in Anadyr Bay

and extended southward along the Koryak coast to Cape Olyutorskiy in the
24 S.A. Macklin (ed.)

0 . 0 0 

0 . 1 0 

0 . 2 0 

0 . 3 0 

0 . 4 0 

3 - 4 4 - 5 5 - 6 6 - 7 7 - 8 8 - 9 9 - 1 0 

A g e s t a n z a 

Avg. increase (kg)

A v e r a g e 

9 3 

9 7 

V o n B M o d e l 

Figure 11: Eastern Bering Sea pollock average individual growth by year as observed from the fishery. The

VonB model represents expected growth increases from a von Bertalanffy curve.

latter half of the year. During the following winter, however, most of the

1997 year class juveniles were distributed under the ice for about 5 months.

In 1998 (a cold year), the abundance of this year class was not very high,

possibly reflecting the negative influence of this long habitation under the

ice.

Acoustic backscatter images at 38 and 120 kHz were compared for the

years 1994–1997 from a survey in mid-September of each year in the Pribilof

Island region of the Bering Sea. Both fish and plankton can be seen in the

images. The plankton patches were indicated by increased backscatter at

120 kHz over 38 kHz, as predicted from geometric modeling of backscatter

from zooplankters. The comparison showed the 4 years to be quite different.

In particular, 1997 appeared to have extremely low fish biomass, and fish

shoals were small and distributed throughout the water column. Plankton

biomass appeared to be large. In contrast, 1996 had extremely high fish

biomass throughout the water column. Plankton biomass, although possibly

obscured by the high fish density, appeared to be low. The cold years of 1994

and 1995 had intermediate fish biomass, with fish shoals restricted to above

or close to the thermocline throughout the day. Plankton biomass during

the daytime was found near the bottom. The low catches of juvenile pollock

around the Pribilof Islands in 1997 were also substantiated by low catches

in trawl collections during the summer and fall in this area (Wilson and

Brodeur, 1998; Brodeur, unpublished data). Given other reports (Coyle,

Stepanenko) that large numbers of young-of-the-year pollock were found in

other areas of the Bering Sea, it is possible that currents were not favorable
FOCI International Workshop Report 25

that year for juvenile pollock to be transported to the Pribilof Islands, which

generally have been an important pollock nursery area.

Distributions of age-1 walleye pollock were compared from data collected

during NMFS bottom trawl surveys during 1996, 1997, and 1998. Age-1

pollock from hauls with more than 100 kg ha−1 were distributed differently

among these 3 years. During 1996 high concentrations of age-1 fish were

landed from 25 stations, no catch occurred at 179 stations, and the largest

catch was 11,537 kg. During 1997, age-1 fish were landed from 21 stations,

no catch at 189 stations, and the largest catch was 1,214 kg. During 1998,

high numbers of age-1 pollock were landed at only 8 stations, no catch at

84 stations, and the largest catch was 656 kg. It appears that age-1 pollock

have decreased in abundance over this time period and in 1998 were more

spread out with smaller catches over a larger area. This would be predicted

from a smaller cold pool area. Food of age-1 pollock varied from 1996 to

1997, with euphausiids being more important in 1996 and mysid shrimp in

1997. Metridia sp. were taken as prey in 1996 but not in 1997, while crab

megalopae were taken only in 1997. Mysids may indicate feeding closer to a

benthic food source in 1997, although pelagic copepods were the dominant

prey item in both years.

The major crab fisheries in the Bering Sea have also shown dramatic

fluctuations over the last several decades. Factors operating during early life

history are thought to be most important to determining year class success.

So, the potential effects of ocean conditions during 1997 and 1998 will not be

realized until those year classes are recruited to the surveyed stocks 6–8 years

in the future. It is too early to predict what the effects of the recent anomalous conditions might have been on crab recruitment, but studies of stock

and environmental trends (Zheng and Kruse, in press) and detailed analyses

of recruitment processes for select individual stocks (Rosenkranz et al., 1998)

shed some light. Long-term trends tend to indicate that some Alaskan crab

stocks show poorer recruitment during periods of strong Aleutian lows and

warmer sea surface temperatures. This is opposite to the relationship shown

for salmon, herring, and most groundfish (Zheng and Kruse, in press). This

pattern appears to hold for Bristol Bay red king crabs. Nearly all Bering

Sea king crabs and Tanner crabs showed consistent downward trends in year

class strength during the early 1970s. However, Bering Sea crab trends since

the late 1970s have been very stock specific. Bristol Bay red king crab recruitment continued to steadily decline until a relatively strong 1990 year

class (consistent with a strong Aleutian low) and appears to have since returned to low recruitment levels. Tanner and snow crab year classes in the

early 1980s were strong, but recruitment of Tanner crabs have since declined

sharply while snow crab year classes continued strong through the 1980s.

Two blue king crab stocks experienced increasing recruitment from brood

years in the late 1970s to early 1980s and were relatively stable through the

late 1980s. From these disparate recent stock trends, it is apparent that no

one climate or ocean factor accounts for all the variability in crab recruitment. In the case of eastern Bering Sea Tanner crabs, it is suggested that

higher productivity and favorable wind conditions promoting transport to

offshore nursery areas are both important variables in recruitment success
26 S.A. Macklin (ed.)

(Rosenkranz et al., 1998). Mild temperatures experienced by maturing adult

Tanner crabs also seem to favor stronger year classes through an effect on

reproduction.

In summary, it is too early to fully assess the impacts of the anomalous

oceanographic conditions during 1997 and 1998 on groundfish recruitment

and other aspects of their life history. Future research needs include:

• additional sampling for pre-recruit stages and forage fishes to track

their abundance trends and map their habitat

• more information on maturity, age, and weight-at-age

• identification of climate conditions linked to recruitment and growth

of groundfish.
FOCI International Workshop Report 27

Seabirds and Marine Mammals

Contributors: C.T. Tynan, G.L. Hunt, Jr., E.H. Sinclair, and C. Pungowiyi

Seabirds and Marine Mammals

The extreme weather events of 1997 and 1998 influenced the survival and

trophic ecology of marine birds in the southeastern Bering Sea by changing

the distribution, abundance, and availability of prey resources. For the period 1997 and 1998, three sources of information on the ecology of marine

birds in the southeastern Bering Sea were available. Vern Byrd of the U.S.

Fish and Wildlife Service, Alaska National Maritime Refuge monitoring program provided information on the reproductive ecology of seabirds nesting

on the Pribilof Islands. SEBSCC and Inner Front projects supported study

of prolonged production, trophic transfer, and processes at Bering Sea inner

fronts. Finally, Vivian Mendenhall of the U.S. Fish and Wildlife Service,

Office of Migratory Bird Management, assembled reports of dead seabirds

observed floating or washed ashore along the coasts of Alaska and the Russian far east.

On the Pribilof Islands, the U.S. Fish and Wildlife Service obtained data

on the reproductive success of nesting kittiwakes (Rissa tridactyla, and R.

brevirostris) and murres (Uria lomvia, and U. aalge). Kittiwakes are surfaceforaging members of the gull family and are limited to obtaining food from

the upper 25 cm of the water column. Murres, in contrast, are pursuit-diving

members of the auk family, and can dive to almost 200 m. Thus murres are

able to reach prey at almost any depth on the southeastern Bering Sea

shelf. In general on the Pribilof Islands, the two species of murres show a

higher average rate of reproductive success and less interannual variation in

reproductive success than the two species of kittiwakes. This is presumably

because kittiwakes are sensitive to the vertical distribution of prey in the

water column, whereas the deep diving murres can access the entire water

column and can seek alternative prey at depth if preferred prey are not

available.

During both 1997 and 1998, the Inner Front project obtained data on

the distribution, abundance, mortality, and trophic ecology of short-tailed

shearwaters (Puffinus tenuirostris) over the middle shelf and coastal domains

of the southeastern Bering Sea, particularly in the vicinity of the inner front.

Short-tailed shearwaters nest in southeastern Australia and migrate to the

Bering Sea, where they forage from May until October. In the Northern

Hemisphere, these birds take a wide variety of prey. However, in the shelf

regions of the southeastern Bering Sea and Bristol Bay, previous studies

have shown that between 80% and 100% of their diet has consisted of adult

euphausiids which they capture in the upper water column to depths of as

much as 35 m.

At the Pribilof Islands in both 1997 and 1998, kittiwake reproductive

success was at or above the long-term mean, whereas murre reproductive

success was considerably below the mean. This unexpected finding may

have been influenced by the presence of the dense coccolithophore bloom
28 S.A. Macklin (ed.)

near the islands. These minute plant organisms are covered with calcium

carbonate plates and reflect light. Objects close to the surface are highlighted by this bright background. Bioluminescent organisms at the surface,

such as lantern fish eaten by kittiwakes, would be conspicuous at night if

their flashes were reflected by the coccolithophores beneath them. Thus the

surface foraging of the kittiwakes may have been enhanced by the presence

of the coccolithophore bloom. In contrast, within and below the coccolithophore bloom, ambient light levels were greatly reduced and scattering of

light was enhanced. These effects of the bloom would decrease the ability

of murres to see and pursue their prey deep in the water. Thus, the poor

reproductive performance of murres nesting on the Pribilof Islands in 1997

and 1998 may have been caused by a decreased ability of murres to obtain

prey at depth.

In June 1997, short-tailed shearwaters were fat and were feeding on

small near-surface patches of adult euphausiids, despite a general lack of

widespread, epibenthic layers of euphausiids in the coastal domain. In late

August and early September 1997, shearwaters were, for the most part, emaciated and often seemed lethargic in their reactions to the approach of the

research vessel. Their diets included juvenile euphausiids, as well as squid

and fish, an unexpectedly broad diet that suggested that the more usually

taken adult euphausiids were unavailable in the shallow waters of the coastal

domain.

During summer and fall 1997, unusually high numbers of seabirds, most

notably short-tailed shearwaters, died in the eastern Bering Sea and in parts

of the Gulf of Alaska. Starting in early August, unusually high numbers

of shearwaters washed ashore along the south side of the Alaska Peninsula,

along the shores of the southeastern Bering Sea and Bristol Bay, on at least

some of the Aleutian Islands, and in the northern Bering Sea at St. Lawrence

Island, and along portions of the Chukotskiy Peninsula of Russia. Within our

study areas, about 163 carcasses were seen floating, and on the basis of the

number seen per unit area searched, we estimate that about 191,500 dead

shearwaters were afloat during our surveys, about 10% of all shearwaters

present in our study areas. Starvation was the apparent cause of mortality.

Many of the shearwaters that we collected were underweight, lacked lipid

reserves, and had lost a significant proportion of their flight muscle mass.

The die-off of shearwaters in 1997 was more widespread than the coccolithophore bloom. Thus, although the dense bloom may have impaired

shearwater foraging, overall some other factor(s) must have influenced the

availability of prey. In 1997, an early spring bloom followed by a secondary

utilization of nutrients below the pycnocline resulted in a shutdown of new

production by the end of May. This lasted until storms in fall resulted in

vertical mixing of nutrients. Consequently, there would have been little new

production to support grazing by euphausiids. Additionally, the exceptionally warm surface temperatures found throughout the eastern North Pacific

Ocean in 1997 may have influenced the distribution of euphausiids. Elsewhere, euphausiids have been found to cease surface swarming when water

temperatures were excessive. Either of the above factors may have resulted

in the euphausiids’ retreat to the deeper waters of the middle shelf domain,
FOCI International Workshop Report 29

where they were present in dense, epibenthic layers and unavailable to shearwaters.

In June 1998, shearwaters were of lighter mass than they were in June

of 1997, apparently because euphausiids, present in extensive, near-bottom

layers in the coastal domain, were late in beginning their mating swarms,

and few near-surface patches were available to the shearwaters. By August

and September 1998, shearwaters exhibited lower body masses than they did

in 1997. Nevertheless, there was no evidence of unusual levels of mortality

in 1998.

How lightweight birds in 1998 avoided starvation is not clear. In the

eastern Bering Sea in 1998, sea surface temperatures were cooler and spring

production was more prolonged than in 1997. This prolonged production

supported euphausiids and the prey of juvenile fish through at least June.

In fall 1998, shearwaters collected in the southeastern Bering Sea had more

fish and squid in their stomachs than was true in past seasons. In particular, in the middle shelf domain and near the inner front southwest of Cape

Newenham, shearwaters fed on age-0 walleye pollock that were present in

great abundance. It is possible that this switch to juvenile fish provided sufficient nutrition for shearwaters to avoid starvation, but insufficient energy

density to permit weight gain.

An additional factor that may be influencing the availability of euphausiids to shearwaters is the possibility of competition with baleen whales. In

the late 1980s near the Pribilof Islands, and in both 1997 and 1998 in the

coastal and middle shelf domains, whales were observed foraging in the vicinity of feeding shearwaters. In most of these instances, the shearwaters were

taking euphausiids, although in one case their prey was juvenile pollock.

C. Tynan (personal communication) has also reported aggregations of large

whales foraging in the vicinity of feeding shearwaters. It is likely that since

the 1970s there has been an increase in the number of whales on the Bering

Sea shelf. The impact of their foraging on the availability of food to other

predators in this ecosystem has yet to be evaluated.

Cetaceans

Coastal and oceanic warming associated with El Ni ̃no can produce redistributions of many marine species, including cetaceans. During the summer of

1997, Pacific white-sided dolphins, a temperate species generally confined to

the Gulf of Alaska and regions further south, were observed in outer Bristol

Bay. Although there have been sightings of this species in other years in the

Bering Sea, their presence in 1997 may have been linked to the warm water

anomalies. Redistributions of large whales may also have occurred.

Large baleen whales migrate long distances to forage in the productive

waters of polar and subpolar regions during summer. As such, they are important indicators and integrators of the regional productivity and biomass

of their principal zooplankton prey (euphausiids and copepods). In the

Bering Sea, migratory baleen whales have historically tracked the “green

belt” areas of high productivity along the shelf-edge region (Springer et al.,

1996). However, during a July 1997 survey of cetaceans, high densities of
30 S.A. Macklin (ed.)

large baleen whales were observed on the middle shelf domain, in and near

the coccolithophore bloom (Tynan, 1998a). The middle shelf domain now

appears to support the highest baleen whale biomass in the southeastern

Bering Sea (Tynan, 1998b). It is important to determine whether these redistributions reflect the anomalous warming of 1997 and 1998, or whether

the movements of whales signal a much longer-term change in the regional

productivity and trophic structure of the Bering Sea ecosystem.

Northern fur seals, which breed during the summer on the Pribilof Islands, were infected with a fungus, possibly Trycophytum, that caused the

loss of guard hair. Although the condition has been observed in pinniped

populations in more southern regions, such as along the coast of California,

this is the first recorded outbreak of the infection in Bering Sea seals. The

outbreak appears to be associated with the warm water anomalies of 1997

and 1998. Up to 15% of the females appear to have been infected. The

longer-term effects on the thermoregulatory ability of the seals is unknown.
FOCI International Workshop Report 31

Conclusions of Working Groups

Compilers: T. Wyllie-Echeverria, E.D. Cokelet, and S.A. Macklin

During the second day of the workshop, participants divided into two discussion groups (Appendix C) to explore three questions that served as themes

for the workshop. Drs. James Schumacher and Tina Wyllie-Echeverria

served as facilitator and rapporteur, respectively, of one group, while Drs.

Alan Springer and Edward Cokelet filled similar roles for the other group.

The collective responses of the two groups are presented here on a questionby-question basis.

1. What are the physical, chemical, and biological mechanisms

and interactions causing the changes in the ecosystem observed in

1997 and 1998?

Climate change in the Bering Sea is difficult to detect because of large

year-to-year variability that can mask a more gradual trend. Nevertheless,

it seems that the meteorological conditions of the winter and spring-summer

seasons have changed between the 1980s and the late 1990s in Alaska. For

1977 through 1989, there was a major warming with a strong Aleutian low

directing warm air from the Pacific toward Alaska. In the 1990s, the low

shifted westward, and high pressure set up over the state circulating cold

arctic air southward over the Bering Sea. Winters have become slightly

colder with more ice, and summers warmer with clearer skies. There has

also been a trend toward more frequent El Ni ̃nos in the 1990s.

Three conditions reinforced one another in 1997 and 1998: (1) a persistent positive North Pacific weather pattern anomaly with associated high

pressure over Alaska and low over Siberia, (2) an El Ni ̃no-reinforced high

pressure over Alaska beginning in April 1997—somewhat earlier than usual

for El Ni ̃no onset, and (3) strong blocking ridges in May of both years. However, there were also differences between the two years. The spring of 1997

was benign with lower than average winds and sunnier days; whereas spring

1998 had stormy weather.

The physical oceanic conditions on the southeastern Bering Sea continental shelf differed between 1997 and 1998. Southward ice extent was above

average in 1997 and below average in 1998. Both summers had warmerthan-average temperatures, but for different reasons. In 1997 the water

was warmed by increased sunshine and lack of wind mixing. In the intervening winter, cold water at the freezing point did not mix to the bottom

as normally occurs, and spring 1998 commenced with warmer water that

heated to higher temperatures in late summer. Perhaps this carry-over of

warm bottom water from the previous summer is a mechanism for climate

change. Spring 1997 winds forced only weak onshelf flow, as verified by

satellite-tracked drifting buoys that showed sluggish onshore and alongshore

flow. Spring 1998 brought strong onshelf flow in the upper few meters of the

water column and strong northward flow of 15–20 cm s−1 along the 50-m

isobath where 3–5 cm s−1 is the norm. This pushed warm water northward,

reinforcing the warm start from the preceding winter. Kamchatka Strait
32 S.A. Macklin (ed.)

is the primary outlet from the Bering Sea. The transport (5.5 Sv) of the

Kamchatka Current was only about half its usual magnitude in 1998.

In the summers of 1997 and 1998, the nutrient reservoir below the pycnocline on the southeastern Bering Sea shelf was depleted, contrary to the

PROBES observations of the 1970s. In 1997, the retreating sea ice was

followed by a phytoplankton bloom that stripped the nutrients from the surface layer. A strong May storm mixed the shelf to 50 m, providing a second

bloom. Then weak winds and increased sunlight caused a shallow mixed

layer below which new phytoplankton growth occurred, stripping the nutrients from the bottom layer. In 1998, the ice may have retreated too early to

trigger a bloom. Stratification remained weak well into June due to storms,

and net phytoplankton growth was low and balanced by zooplankton grazing. Net new production continued over a longer period of time, ultimately

consuming the nutrients in the entire water column.

Primary production in 1997 and 1998 may have been higher than normal

on the southeastern shelf because the entire nutrient pool was utilized. Sediment trap observations of the carbon flux owing to new production during

1997 were high (∼300 mg C m−2 d−1), thus supporting the conjecture.

Unusual coccolithophore blooms covered large areas of the shelf both

years as shown by chalky, aquamarine water in SeaWIFS satellite realcolor images. These calcium-carbonate-rich phytoplankton are favored over

silicate-based diatoms when ammonium concentrations are high (due to

zooplankton grazing and nutrient recycling) and silicate concentrations are

low due to depletion by diatoms. Water temperatures above 10◦C, as observed, favor the coccolithophore E. huxleyi over Phaeocystis, another oftendominant phytoplankter. Preliminary sediment trap analyses from a single

location do not show coccoliths in zooplankton fecal pellets, implying that

they were not preyed upon. Diatoms were present in the traps, but conclusions about relative abundance remain open. Water samples from October 1997 show that coccolithophores comprised 98% of the phytoplankton

biomass.

Isotopic-ratio measurements in whale baleen indicate a long-term drop

in primary production near Bering Strait. D. Schell hypothesizes that reduced flow through the strait leads to less onshelf flow, lower nutrient flux

(replenishment), and less productivity.

No consistent picture emerges for zooplankton on the southeastern shelf,

but the measurements are sparse. Euphausiid abundance was low at the

inner front in 1997, but very high over the middle shelf domain near the

Pribilof Islands. It is puzzling that oceanic copepods were pushed into the

inner front (∼50 m) in June 1998, but that onshore flow was low that year.

Pollock larvae were located more to the north in 1997 than in the early 1990s,

and their growth rate was low. Larval concentrations were high across the

middle shelf in 1998.

Pollock hydroacoustic and trawl survey data for 1997 were not too different from other years on the shelf (N. Williamson, personal communication).

In September 1997, there were strikingly fewer age-0 pollock in close proximity of the Pribilofs than observed since 1994 (G. Swartzman, personal

communication).
FOCI International Workshop Report 33

Sockeye salmon returns were lower than expected in 1997 and 1998 in

Bristol Bay, but escapement goals were met. Yukon chinook salmon runs

also were low, and the fish were small and late returning. Low runs also were

reported for the southern side of the Alaskan Peninsula and Kodiak Island.

Salmon in some river systems fared better. Russia had unexpectedly high

salmon runs. Returning fish were being stripped of their roe and bulldozed

back into the sea due to processing under-capacity.

Scientists find it difficult to imagine how the numbers of returning adult

salmon could be affected by environmental conditions in the Bering Sea.

The fish returning to Bristol Bay were not emaciated or diseased, although

they were smaller than average. However, it is probable that salmon smolts

heading out to sea would be affected by climatic conditions. Those effects

would be noted in 2 to 3 years when the salmon returned to spawn. Thus,

for salmon returning in 1997 and 1998, perhaps conditions in 1994 to 1996

should be investigated. Similarly, salmon returns in 1999 to 2001 may be

affected by the recent anomalies.

We do not yet know enough about the ecosystem and its workings to

surmise any conclusions about the co-occurrence of coccolithophore blooms,

bird die-offs, low salmon runs, increased whale sightings, etc.

2. What research is needed to better understand the origin and

persistence of these changes?

Participants felt that both top-down and bottom-up approaches are useful in understanding the Bering Sea ecosystem. It is helpful to begin with atmospheric forcing and its affects on the ocean environment, then to consider

the consequences of the basic biology of animals. The key to understanding broad-scale changes in this ecosystem seems to lie in our understanding

of temperature, winds, and sea ice variability. Monitoring programs incorporating moored and drifting instruments and knowledge gained from local

users of the Bering Sea combined with retrospective studies help direct future

questions and place the present situation in a long-term context. Multidisciplinary, process-oriented studies are necessary and should focus on specific

questions of a regional nature to attain understanding of interactions among

species and their environment. Emphasis should be placed on knowledge

of individual species’ life histories and how those species interrelate to other

key species. We need to understand E. huxleyi, its role in the carbon budget,

and how it differs from another small cell that is abundant in the summer,

Phaeocystis sp. An iterative process where we return to retrospective investigations using the knowledge gained through process work may provide

clues to key hypotheses. Additionally, modeling may help predict how key

species will respond to hypothesized environmental conditions.

Communication is vital to progress in science. Besides established methods of communication, participants suggested establishing a chat room as

part of the Bering Sea and North Pacific Ocean theme page (http://

www.pmel.noaa.gov/bering/) to facilitate information exchange on a timely

basis.

The recently drafted Science Plan for the Bering Sea Ecosystem (avail-
34 S.A. Macklin (ed.)

able from the theme page at http://www.pmel.noaa.gov/bering/

pages/inter-agency/) was recognized as an accurate and timely guide to

research needs. Specific research recommendations from this workshop include:

• Measure the system over the next 3 years to capture critical evidence

to support or reject claims of a climatological regime shift that may

be occurring in the Bering Sea. This opportunity was missed before,

and may not recur for decades.

• Establish process-oriented research to determine intervening pathways

between atmospheric forcing and ecological response.

• Monitor ocean circulation and water properties at more locations for

longer time periods to establish baseline values. Biophysical drifters

and moorings are needed. Additional sites in the outer shelf and

coastal domains between the Pribilof Islands and St. Matthew Island

are needed to expand the present FOCI measurement area.

• Provide more observations of the flow through the Aleutian Islands’

passes to determine the balance of heat and salt in the Bering Sea

basin.

• Consider the flow through Kamchatka and Bering Straits. The oceanic

flow into the Bering Sea enters through a variety of passes, but exits

through just these two constrictions. They would make natural pulse

points to measure the effect of climate on the mean flow field over long

time periods.

• Monitor Bering Strait flow and its connection with onshelf nutrient

flux and primary production in the region.

• Couple physical measurements with those of primary and secondary

productivity.

• Determine the specific fate of primary production. How do variations

in biomass, production, and species composition affect the transfer of

energy to apex predators?

• Investigate whether recently observed changes are propagating through

the entire trophic pyramid.

• Increase the sampling frequency. Provide more cruises to capture the

spring and summer production pulses and food web dynamics. Shorebased sampling from smaller vessels in the Pribilofs and along the

northern Alaskan Peninsula should be considered. A mosaic of regions

and ecological steps should be constructed and sampled at key points

and times.

• Study predator-prey relationships. They are important to the ecosystem at all levels.
FOCI International Workshop Report 35

• Research connections between the pelagic and benthic communities.

• Learn more about forage fish (myctophids, capelin, eulachon, sand

lance) and squid.

• Determine the reason(s) for dramatic increases in jellyfish population.

• Monitor salmon and environmental conditions along the northern side

of the Alaskan Peninsula in late summer and early fall.

• Establish when year-class strength is set for different Bering Sea salmon

stocks. Could it be when smolts migrate along the Alaskan Peninsula

in May to August?

• Investigate the relationship between salmon in the eastern, central,

and western Bering Sea.

• Study the movement of juvenile pollock on the basin scale.

3. Given the large changes in the Bering Sea ecosystem, how do

we manage marine resources in a fluctuating and highly variable

environment?

Managers need a broad, simple rule by which to manage. It is the researchers’ challenge to understand species responses in a variable environment and be able to predict large-scale consequences. Because species have

evolved in this variable environment, it was suggested that human management of its resources also be flexible. Harvesters need to be able to switch

between targeted species not only as seasons vary, but as populations vary

interannually. This would require a more flexible, integrative management

policy. At the governmental level, official policies and mechanisms should be

established to exchange scientific data and manage the international pollock

fishery (Russia, USA, donut hole, etc.).

Recommendations

Recent events have demonstrated how quickly and dramatically the Bering

Sea ecosystem responds to changes in established weather patterns. These

patterns are the result of decadal and shorter-term variations in the atmosphere. Participants of the workshop emphasized the scientific and socioeconomic value of the Bering Sea ecosystem. The recommendations below seek a more complete understanding of the ecosystem so that potential

changes may be forecast and distributed to all stakeholders (e.g., as is done

with El Ni ̃no forecasting). The recommendations are consistent with those

in the recently published Draft Science Plan for the Bering Sea Ecosystem

sponsored by NOAA, Alaska Department of Fish and Game, and the U.S.

Department of the Interior.

• Design and immediately implement a long-term ecosystem observing

system to continually document the state of the system and observe future changes. System components would include information obtained

from scientific observing systems, industry, and Native Alaskans.
36 S.A. Macklin (ed.)

• Begin a comprehensive synthesis of our knowledge of the ecosystem.

Components would include paleo-oceanographic data, anthropological

data, results of prior research projects, and traditional knowledge.

• Assemble integrated teams of scientists and stakeholders to determine

how the Bering Sea ecosystem operates. Teams would conduct processoriented research to determine how changes in physical forcing, biological production, biomass, and species composition affect the transfer

of carbon from lower trophic levels to living marine resources and protected species.

• Design, assemble, and test analytical, numerical, and conceptual models that seek to examine critically our knowledge of how the system

operates. The long-term goal for the models is accurate forecasting of

system response to physical and biological perturbation.

• Build tools (and collaborative relationships) that facilitate communication among ecosystem stakeholders (industry, Native Alaskans, representatives of state and federal agencies).

• Design and implement flexible resource management strategies that

can quickly interpret and incorporate relevant information about resources and changing conditions into management decisions.

Acknowledgments. The conveners and organizers gratefully acknowledge the

North Pacific Fishery Management Council, the Alaska Fisheries Science Center,

and the Pacific Marine Environmental Laboratory for their support for this workshop. We are especially appreciative of the keynote speakers who established the

basis of understanding that allowed workshop participants to discuss together recent conditions in the Bering Sea. Thanks, too, to the international and voluntary

speakers who augmented our understanding. Mr. Bart Eaton, Trident Seafoods

Corporation, reminded us of partnerships that industry and science can build. Discussion group facilitators and rapporteurs skillfully guided and recorded discussions.

Finally, a truly heartfelt thanks to Lynn Long, Kurt Zegowitz, Joyce Gearhart,

Destry Wion, Sonia Hamilton, Chris Baier, and Jen Quan who created the ambience for an enjoyable and productive workshop.

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observations of water column temperature, salinity and spring phytoplankton

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38 S.A. Macklin (ed.)

Appendix A: Attendees

Dr. Knut Aagaard

Applied Physics Laboratory

P.O. Box 355640

University of Washington

Seattle, WA 98195-5640

USA

Phone: 206-543-8942

E-mail: aagaard@apl.washington.edu

Dr. Milo Adkinson

University of Alaska Fairbanks—Juneau Center

School of Fisheries and Ocean Sciences

1120 Glacier Highway

Juneau, AK 99801

USA

Phone: 907-465-6251

E-mail: ffmda@uaf.edu

Ms. Cheryl Baduini

Department of Ecology and Environmental Biology

University of California, Irvine

Irvine, CA 92697-2525

USA

Phone: 949-824-4747

E-mail: cbaduini@uci.edu

Ms. Chris Baier

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4112

E-mail: cbaier@afsc.noaa.gov

Dr. Kevin Bailey

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4243

E-mail: kbailey@afsc.noaa.gov

Dr. Eddie Bernard

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6800

E-mail: bernard@pmel.noaa.gov

Dr. Nick Bond

JISAO at NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6459

E-mail: bond@pmel.noaa.gov

CAPT Dale Bretschneider

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6813

E-mail: bret@pmel.noaa.gov

Dr. Ric Brodeur

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4318

E-mail: rbrodeur@afsc.noaa.gov

Dr. G. Vernon Byrd

Alaska Maritime National Wildlife Refuge

2355 Kachemak Bay Drive, Suite 101

Homer, AK 99603

USA

Phone: 907-235-6546

E-mail: vernon byrd@fws.gov

Ms. Molly Chythlook

P.O. Box 692

Dillingham, AK 99576

USA

Phone: 907-842-5925

E-mail: mollyc@fishgame.state.ak.us

Dr. James Coe

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4000

E-mail: jim.coe@noaa.gov

Dr. Ned Cokelet

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6820

E-mail: cokelet@pmel.noaa.gov

Dr. Ken Coyle

Institute of Marine Science

University of Alaska Fairbanks

Fairbanks, AK 99775-7220

USA

Phone: 907-474-7705

E-mail: coyle@ims.alaska.edu
FOCI International Workshop Report 39

Dr. Ned Cyr

Office of Science and Technology

NOAA/ST2 Room 14348

1315 East-West Highway

Silver Spring, MD 20910

USA

Phone: 301-713-2367

E-mail: ned.cyr@noaa.gov

Mr. Bart Eaton

Trident Seafoods Corporation

5303 Shilshole Ave. NW

Seattle, WA 98107

USA

Phone: 206-783-3818

E-mail: cassieowens@tridentseafoods.com

Dr. Mikhail Flint

Department of Plankton Ecology

P.P. Shirshov Institute of Oceanology RAS

36 Nakhimovski Prospekt

Moscow, 117851

Russia

Phone: 7-095-124-77-49

E-mail: flinteco@glas.apc.org

Dr. Dave Fluharty

School of Marine Affairs

University of Washington

Box 355685

Seattle, WA 98195-5685

USA

Phone: 206-685-2518

E-mail: fluharty@u.washington.edu

Dr. Toni Frohoff

The Humane Society of the United States

321 High School Road #374

Bainbridge Is., WA 98110

USA

Phone: 206-780-2532

E-mail: frohoff@ix.netcom.com

Dr. Kathy Frost

Alaska Department of Fish and Game

1300 College Road

Fairbanks, AK 99701

USA

Phone: 907-459-7214

E-mail: kfrost@fishgame.state.ak.us

Dr. John Goering

Institute of Marine Science

University of Alaska Fairbanks

Fairbanks, AK 99775-7220

USA

Phone: 907-474-7895

E-mail: goering@ims.alaska.edu

Ms. Sonia Hamilton

JISAO at NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6746

E-mail: sham@pmel.noaa.gov

Dr. Steve Hare

International Pacific Halibut Commission

University of Washington

Box 95009

Seattle, WA 98195-7980

USA

Phone: 206-634-1838

E-mail: hare@iphc.washington.edu

Dr. Jack Helle

NMFS/AFSCF/AKC41

1305 Glacier Highway

Juneau, AK 99801-8626

USA

Phone: 907-789-6038

E-mail: Jack.Helle@noaa.gov

Dr. Susan Henrichs

Institute of Marine Science

University of Alaska Fairbanks

Fairbanks, AK 99775-7220

USA

Phone: 907-474-7807

E-mail: henrichs@ims.alaska.edu

Dr. Al Hermann

JISAO at NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6495

E-mail: hermann@pmel.noaa.gov

Dr. Anne Hollowed

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4223

E-mail: hollowed@afsc.noaa.gov

Dr. George Hunt

Department of Ecology and Environmental Biology

321 Steinhaus Hall

University of California Irvine

Irvine, CA 92697-2525

USA

Phone: 714-824-6322

E-mail: glhunt@uci.edu
40 S.A. Macklin (ed.)

Dr. Jim Ianelli

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6510

E-mail: Jim.Ianelli@noaa.gov

Dr. Nancy Kachel

JISAO at NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6780

E-mail: nkachel@pmel.noaa.gov

Dr. Art Kendall

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4108

E-mail: kendall@afsc.noaa.gov

CDR Mark Koehn

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6197

E-mail: koehn@pmel.noaa.gov

Dr. Gordon Kruse

Alaska Department of Fish and Game

Division of Commercial Fisheries

Box 25526 or 1255 W. 8th St.

Juneau, AK 99802-5526

USA

Phone: 907-465-6106

E-mail: gordonk@fishgame.state.ak.us

LCDR Mike Lemon

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6804

E-mail: lemon@pmel.noaa.gov

Ms. Pat Livingston

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4242

E-mail: pliving@afsc.noaa.gov

Dr. Loh-Lee Low

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4190

E-mail: loh-lee.low@noaa.gov

Mr. Allen Macklin

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6798

E-mail: macklin@pmel.noaa.gov

Dr. Nate Mantua

JISAO

University of Washington

Box 354235

Seattle, WA 98195-4235

USA

Phone: 206-616-5347

E-mail: mantua@atmos.washington.edu

Dr. Ole A. Mathiesen

University of Alaska Fairbanks—Juneau Center

School of Fisheries and Ocean Sciences

1120 Glacier Highway

Juneau, AK 99801

USA

Phone: 907-465-6442

E-mail: ffoam@uaf.edu

Dr. Dennis Moore

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4146

E-mail: dmoore@pmel.noaa.gov

Dr. Susan Moore

NMFS/NMML

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4021

E-mail: sue.moore@noaa.gov

Dr. Calvin Mordy

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6870

E-mail: mordy@pmel.noaa.gov
FOCI International Workshop Report 41

Dr. Jeff Napp

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4148

E-mail: Jeff.Napp@noaa.gov

Dr. Jim Overland

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6795

E-mail: overland@pmel.noaa.gov

Mr. Caleb Pungowiyi

Kawerak, Inc.

P.O Box 948

Nome, AK 99762

USA

Phone: 907-443-4380

E-mail: calebp@nome.net

Mr. Ron Reed

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6818

E-mail: reed@pmel.noaa.gov

Ms. Sigrid Salo

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-6802

E-mail: salo@pmel.noaa.gov

Dr. Don Schell

Institute of Marine Science

University of Alaska Fairbanks

Fairbanks, AK 99775-7220

USA

Phone: 907-474-7211

E-mail: schell@ims.alaska.edu

Dr. Jim Schumacher

Two Crow Environmental Consultants

P.O. Box 215

Silver City, NM 88062

USA

Phone: 505-388-2742

E-mail: jdschu@pmel.noaa.gov

Dr. Naonubo Shiga

Faculty of Fisheries

Hokkaido University

3-1-1, Minato-cho

Hakodate, Hokkaido 041-8611

Japan

Phone:

E-mail: nao@pop.fish.hokudai.ac.jp

Ms. Beth Sinclair

NOAA/NMML

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6466

E-mail: beth.sinclair@noaa.gov

Dr. Dave Somerton

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4116

E-mail: David.Somerton@noaa.gov

Ms. Suzanne Speckman

Applied Physics Laboratory

University of Washington

Box 355640

Seattle, WA 98105-6698

USA

Phone: 206-545-8107

E-mail: speckman@fish.washington.edu

Dr. Paul Spencer

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4248

E-mail: paul.spencer@noaa.gov

Dr. Alan Springer

Institute of Marine Science

University of Alaska Fairbanks

Fairbanks, AK 99775-7220

USA

Phone: 907-474-6213

E-mail: ams@ims.uaf.edu

Dr. Phyllis Stabeno

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6453

E-mail: stabeno@pmel.noaa.gov
42 S.A. Macklin (ed.)

Dr. Gary Stauffer

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4170

E-mail: staufferg@afsc.noaa.gov

Dr. Mikhail Stepanenko

TINRO/PRIFO4

Shevchenko Alley

Vladivostok, 690600

Russia

Phone:

E-mail: root@tinro.marine.su

Dr. Dean A. Stockwell

School of Fisheries and Ocean Sciences

P.O. Box 757220

University of Alaska Fairbanks

Fairbanks, AK 99775-7220

USA

Phone: 907-474-5556

E-mail: dean@ims.uaf.edu

Dr. Gordie Swartzman

Applied Physics Laboratory

University of Washington

Box 355640

Seattle, WA 98105-6698

USA

Phone: 206-543-0061

E-mail: gordie@apl.washington.edu

Ms. Carol Tocco

National Marine Fisheries Service

Alaska Region, Public Affairs

P.O. Box 21668 or 709 West 9th Street

Juneau, AK 99802-1668

USA

Phone: 907-586-7032

E-mail: Carol.Tocco@noaa.gov

Dr. Cynthia Tynan

JISAO at NMFS/NMML

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4033

E-mail: tynan@afsc.noaa.gov

Ms. Tiffany Vance

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6767

E-mail: vance@pmel.noaa.gov

Dr. Diana E. Varela

Dept. of Earth and Ocean Sciences

Oceanography, 6270 University Boulevard, Room 1461

University of British Columbia

Vancouver, BC V6T 1Z4

Canada

Phone: 604-822-5517

E-mail: varela@unixg.ubc.ca

Mr. Trey Walker

Fisheries Research Institute

University of Washington

Box 357980

Seattle, WA 98195-7980

USA

Phone: 206-543-7281

E-mail: rwalker@fish.washington.edu

Dr. Bernard Walter

Northwest Research Associates

P.O. Box 3027

Bellevue, WA 98009

USA

Phone: 425-644-9660 ext. 320

E-mail: walter@nwra.com

Mr. Gary Walters

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4143

E-mail: Gary.Walters@noaa.gov

Mr. Brad Warren

Editor, Pacific Fishing Magazine

1515 NW 51st St.

Seattle, WA 98107

USA

Phone: 206-789-5333

E-mail: brad@salmonbay.com

Dr. Gunter Weller

CIFAR

Room 506C Elvey Bldg.

University of Alaska Fairbanks

Fairbanks, AK 99775-7748

USA

Phone: 907-474-6722

E-mail: gunter@gi.alaska.edu

Dr. Terry Whitledge

Institute of Marine Science

University of Alaska Fairbanks

P.O. Box 757220

Fairbanks, AK 99775-7220

USA

Phone: 907-474-7229

E-mail: terry@ims.uaf.edu
FOCI International Workshop Report 43

Mr. Neal Williamson

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6417

E-mail: nealw@afsc.noaa.gov

Dr. Chris Wilson

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6435

E-mail: cdwilson@afsc.noaa.gov

Mr. Matt Wilson

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6522

E-mail: mwilson@afsc.noaa.gov

Mr. Destry Wion

NMFS/AFSC

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-4605

E-mail: destry.wion@afsc.noaa.gov

Dr. Warren Wooster

School of Marine Affairs

University of Washington

Box 355685

Seattle, WA 98195-5685

USA

Phone: 206-685-2497

E-mail: wooster@u.washington.edu

Mr. Bruce A. Wright

NOAA/NMFS11305 Glacier Highway

Juneau, AK 99801

USA

Phone: 907-789-6601

E-mail: bruce.wright@noaa.gov

Dr. Tina Wyllie-Echeverria

1027 Fir Ave.

Provo, UT 84604

USA

Phone: 512-749-6769/801-356-3118

E-mail: tinawe@pmel.noaa.gov

LTJG Kurt Zegowitz

NOAA/PMEL

7600 Sand Point Way N.E.

Seattle, WA 98115-0070

USA

Phone: 206-526-6809

E-mail: zegowitz@pmel.noaa.gov
44 S.A. Macklin (ed.)

Appendix B: Agenda

Monday, November 9, 1998

9:00 am Welcome Dr. Arthur Kendall, Jr. and

Dr. Phyllis Stabeno

9:15 am Pacific climate in 1997–98: El Ni ̃no? PDO?

or another animal from the climate zoo?

Dr. Nathan Mantua

University of Washington

9:45 am Temporal variability of water properties and

currents over the eastern Bering Sea shelf

Dr. Phyllis Stabeno

NOAA/Pacific Marine Environmental Laboratory

10:15 am Bering Sea nutrient dynamics: Then and now Dr. Terry Whitledge

University of Alaska Fairbanks

10:45 am Break

11:15 am Southeastern Bering Sea zooplankton:

Known patterns and potential anomalies

Dr. Jeffrey Napp

NMFS/Alaska Fisheries Science Center

Dr. Kenneth Coyle

University of Alaska Fairbanks

11:45 am Responses of marine birds to unusual conditions in the southeastern Bering Sea, 1997

and 1998

Dr. George Hunt, Jr.

University of California Irvine

12:15 pm Lunch

1:30 pm Redistributions of cetaceans in the southeastern Bering Sea relative to anomalous oceanographic conditions during the 1997 El Ni ̃noDr. Cynthia Tynan

University of Washington

2:00 pm Implications of recent climate changes on

salmon and crabs in the Bering Sea

Dr. Gordon Kruse

Alaska Department of Fish and Game

2:30 pm Bering Sea groundfish abundance and recruitment trendsMs. Patricia Livingston

NMFS/Alaska Fisheries Science Center

3:00 pm Break

3:20 pm International Perspectives:

Nitrogen and iron effects on the physiology

of a coccolithophore and a diatom from the

oceanic Northeast Pacific

Dr. Diana Varela

University of British Columbia

Coccolithophorids in phytoplankton of the

Pribilof region of the Bering Sea

Dr. Mikhail Flint

Shirshov University

Russian Academy of Sciences

Lower trophic conditions coupling with the

recent warm sea surface temperature in the

southeastern Bering Sea shelf in summer

Dr. Naonubo Shiga

Hokkaido University

Walleye pollock distribution and spawning in

the eastern and western Bering Sea

Dr. Mikhail Stepanenko

TINRO
FOCI International Workshop Report 45

4:00 pm Voluntary Presentations:

Marine mammals Mr. Caleb Pungowiyi

Age-1 pollock Dr. Tina Wyllie-Echeverria

Sediment trap and zooplankton stable isotopic compositionDr. Susan Henrichs

Weather Dr. Jim Overland

Northern fur seal pelage condition Ms. Beth Sinclair

Zooplankton-fish acoustic comparison Dr. Gordie Swartzman

Climate change impacts Dr. Gunter Weller

Coccolithophores in the 1970s and 1980s? Ms. Tiffany Vance

TOPEX/Poseidon-ERS2 sea surface height

anomalies

Dr. Ned Cokelet

5:30 pm Reception

6:00 pm Keynote Address Mr. Bart Eaton, Vice President

Trident Seafoods Corporation

7:00 pm Adjourn
46 S.A. Macklin (ed.)

Tuesday, November 10, 1998

8:10 am Coffee

8:30 am Instructions for Discussion Groups Organizing Committee

8:45 am Discussion:

What are the physical, chemical, and

biological mechanisms and interactions causing the changes in the ecosystem observed in

1997 and 1998?

What research is needed to better understand the origin and persistence of these

changes?

Given the large changes in the Bering

Sea ecosystem, how do we manage marine

resources in a fluctuating and highly variable

environment?

As assigned

11:00 am Break

11:30 am Discussion Group Reports Rapporteurs

12:30 pm Closing Comments

12:45 pm Adjourn
FOCI International Workshop Report 47

Appendix C: Discussion Group Assignments

Group 1 (Oceanographer Room)

Schumacher, J.—Facilitator

Wyllie-Echeverria, T.—Rapporteur

Aagaard, K.

Bernard, E.

Bond, N.

Bretschneider, D.

Chythlook, M.

Cyr, N.

Frost, K.

Goering, J.

Hare, S.

Helle, J.

Hermann, A.

Hollowed, A.

Hunt, G.

Ianelli, J.

Kendall, A.

Low, L-L.

Macklin, A.

Mathieson, O.

Napp, J.

Shiga, N.

Tynan, C.

Varela, D.

Warren, B.

Weller, G.

Whitledge, T.

Group 2 (Rooms A and B)

Springer, A.—Facilitator

Cokelet, N.—Rapporteur

Adkinson, M.

Baduini, C.

Bailey, K.

Brodeur, R.

Byrd, V.

Coyle, K.

Flint, M.

Henrichs, S.

Kruse, G.

Livingston, P.

Mordy, C.

Overland, J.

Pungowiyi, C.

Schell, D.

Sinclair, B.

Speckman, S.

Stabeno, P.

Stepanenko, M.

Stockwell, D.

Swartzman, G.

Tocco, C.

Walker, T.

Walter, B.

Williamson, N.

Wright, B.