Distribution and Ecological Interactions off Forage Fishes in the Northern California Current Richard D. Brodeur Fish Ecology Division Northwest Fisheries Science Center NOAA Fisheries With help from Elizabeth Daly, Bob Emmett, Mary Hunsicker, Todd Miller, John Buchanan, Toby Auth, Greg Krutzikowski, Marisa Litz, and many others

1 Forage fish are key links in food webs

Image: Pew Trusts Talk Objectives – 1) Examine changes in larval abundance of important forage fishes in relation to changing climate 2) Summarize surveys which examine distributions of key forage fishes 3) Examine diets of important forage fishes and diet overlap of other species that potentially compete with these forage fishes 4) Examine important fish predators and estimate consumption of forage fishes by pelagic fishes and salmon Winter (Jan. – March) Ichthyoplankton from Newport Line Samples 3.0 EngraulisAnchovy mordax SardinopsSardine sagax OsmeridaeSmelts ) 2.5 3 AmmodytesSand Lance spp. SebastesRockfish spp. StenobrachiusNorthern Lampfish leucopsarus 2.0 ParophrysEnglish Sole vetulus IsopsettaButter Sole isolepis CitharichthysSanddab spp. 1.5 PsettichthysSand Sole melanostictus

1.0

Concentration (number/1000m 0.5

0.0

1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 Year • Earliest (by three months) and most widespread spawning of anchovies and sardines in NCC • Also found Pacific hake and jack mackerel eggs and larvae off Newport

Auth et al. (2019) Global Change Biology

Nearshore (NH 1-15) Density

) 3

- 5 EngraulisNorthern Anchovy mordax SardinopsPacific Sardine sagax 4

3

2

1 *

mean concentration (no. 1000 m 1000 (no. concentration mean 0

10 Log Month/Year Continued through March 2017! * No samples collected Winter Ichthyoplankton composition PCO axis 1 scores and juvenile salmon success

40

2011 2003 2010 20 2015 2013 2016 2012 1998 2005 2002 2008 2019 0 2006 2001 2017 2018 2004 2009 2000 2014 -20

2007 1999

PCO2 (15.5% of total variation)

-40 Salmon returns higher Salmon returns lower

-40 -20 0 20 40 PCO1 (35.5% of total variation) Juvenile salmon that have migrated into the ocean since the warm blob have all had poor subsequent adult salmon returns to the Columbia River 4 Spring Chinook Fall Chinook 3 Coho OPIH Steelhead Unmarked Steelhead 2

1

0

-1

Normalized SalmonNormalized Return data -2

-3

1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017

Outmimigration Ocean Year Adapted from Daly et al. (2019) Forage Fish Study Sampling Years:1998-2009

Sampled two transects off the Columbia River and Willapa Bay every 10 days from April to early August Average Forage Fish Densities 8000 7000 Northern anchovy

-1 ) Pacific herring 3 6000

m

6 5000 Whitebait smelt Pacific sardine 4000 3000 2000 Number Number (10 1000 0 0-10 10-20 20-30 30-40 40-50 50-60 Distance from shore (km)

Emmett et al. (2006) Prog. Oceanogr. Forage Fish Densities

) 9000

3 Whitebait smelt m

6 8000 Pacific sardine 7000 Pacific herring 6000 5000 Northern anchovy 4000 3000 2000

Average Densities (#/10 Average 1000 0 1998 1999 2000 2001 2002 2003 2004 2005 2006 Year Large-Scale Distribution of Pacific Sardines - June

16 - 25 Jun 98 16 - 24 Jun 99 29 May - 25 Jun 00 24 Jun - 1 Jul 01 1 - 28 Jun 02 23 Jun - 3 Jul 03 22 - 29 Jun 04

48°N 48°N

47°N 47°N

46°N 46°N

e

d

u

t i 45°N 45°N

t

a

L

44°N 44°N

Medium Sized Large Sized Density (no./106 m3) Density (no./106 m3) 43°N 0 to <0.001 0 to <0.00143°N 0.001 to <100 0.001 to <100 100 to <1,000 100 to <1,000 1,000 to <10,000 1,000 to <10,000 42°N 42°N 10,000 to <44,975 10,000 to <44,975

125°W 124°W 125°W 124°W 125°W 124°W 125°W 124°W 125°W 124°W 125°W 124°W 125°W 124°W Longitude Emmett et al. (2006) CalCOFI Reports Large-Scale Distribution of Pacific Sardines - September 20 - 29 Sep 98 21 Sep - 1 Oct 99 19 - 24 Sep 00 21 - 28 Sep 01 26 Sep - 3 Oct 02 26 Sep - 3 Oct 03 22 - 29 Sep 04

48° N 48°N

47° N 47°N

46° N 46°N

e

d

u

t i 45° N 45°N

t

a

L

44° N 44°N

Small Sized Medium Sized Large Sized Density (no./106 m3) Density (no./106 m3) Density (no./106 m3) 43° N 0 to <0.001 0 to <0.001 0 to <0.001 43°N 0.001 to <100 0.001 to <100 0.001 to <100 100 to <1,000 100 to <1,000 100 to <1,000 1,000 to <10,000 1,000 to <10,000 1,000 to <10,000 42° N 42°N 10,000 to <44,975 10,000 to <44,975 10,000 to <44,975

125°W 124°W 125° W 124° W 125°W 124°W 125°W 124°W 125°W 124°W 125°W 124°W 125°W 124°W Longitude

Emmett et al. (2006) CalCOFI Reports Predicted Probabilities of Occurrence in June

49° 49° 49° 49°

48° 48° 48° 48°

47° 47° 47° 47°

46° 46° 46° 46°

45° 45° 45° 45°

Surf Smelt Herring Anchovy Sardine 44° 44° 44° 44°

125.5° 124.5° 123.5° 125.5° 124.5° 123.5° 125.5° 124.5° 123.5° 125.5° 124.5° 123.5°

Fitted simple generalized additive models (GAMs) to each species separately. The response variable was presence/absence of a species at the selected stations and the explanatory variables included start lat, start lon, bottom depth and year. Using the models, calculated the predicted probabilities of occurrence of each species across the sampling area.

Provided by Mary Hunsicker Interannual Variability in Diets of Forage Fishes

Northern anchovy (Engraulis mordax) Pacific herring (Clupea pallasii)

Pacific sardine (Sardinops sagax) Surf smelt (Hypomesus pretiosus)

Jack mackerel (Trachurus symmetricus) Whitebait smelt (Allosmerus elongatus) Diet composition in June by weight

Copepods

Gelatinous Zooplankton

Euphausiids

A C C W W A A C C W W A A C C W W

Brodeur et al. (2019) Marine Ecology Progress Series Diet composition in June by weight

A A C C W W A A C W W A A W W

Brodeur et al. (2019) Marine Ecology Progress Series Forage Fish Condition Between Ocean Regimes

Fatter Condition

Leaner

Brodeur et al. (2019) Marine Ecology Progress Series AVERAGE WARM

High

Trophic Level

Low

18 Hunsicker et al. (In prep) Pelagic and demersal fish predators on juvenile and adult forage fishes in the Northern California Current: Spatial and Temporal Variations

Richard Brodeur1, John Buchanan1, and Robert Emmett2

1Northwest Fisheries Science Center, NOAA Fisheries, Newport, OR

2Northwest Fisheries Science Center, NOAA Fisheries, Hammond, OR

CalCOFI Reports 2014 Methods

Conducted literature survey of diet studies along the coast of Washington, Oregon, and northern California.

• Multispecies surveys of diets • Studies on individual species or related species in small geographic area • Published and unpublished studies (theses) • Potential pelagic, midwater, and demersal fish predators • Summarized forage prey by percent of total weight in diet

Brodeur, Buchanan, and Emmett (2014) CalCOFI Rep. Forage Fishes Examined

Pacific saury (Cololabis saira) Pacific herring (Clupea pallasii)

Pacific sardine (Sardinops sagax) Juvenile hake (Merluccius productus)

Northern anchovy (Engraulis mordax)

Juvenile rockfish (Sebastes spp.)

Smelt (Osmeridae) Predator Fishes Examined

Elasmobranchs Demersal fishes • Spiny dogfish • Lingcod • Blue shark • Black rockfish • Soupfin shark • Yellowtail rockfish • Bocaccio Pelagic fishes • Yelloweye rockfish • Jack mackerel • Rougheye rockfish • Pacific hake √ • Sablefish • Chinook salmon √ • Pacific halibut • Coho salmon • Arrowtooth flounder • Albacore

Brodeur, Buchanan, and Emmett (2014) CalCOFI Rep. Hake Diets from West Coast Bottom Trawl Surveys West Coast Hake Diets by Size Hake in WC Groundfish Surveys N = 120 97 118 93 27 N = 996 131 74 100 100

80 80 Euphausiids Crustaceans Cephalopods Euphausiids Anchovy Crustaceans Smelt Herring 60 60 Rockfish Anchovy Herring Smelt Hake Rockfish Other Fishes Hake 40 Other Fishes 40

Percent weight contributionPercent weight

Percent weight contribution weight Percent

20 20

0 0

35-39 40-44 45-49 50-54 > 55

Summer 1989Autumn 1991Autumn 1992 Rexstad and Pikitch (1986) Buckley et al. (1999) Adult Chinook Diets off Oregon and Washington

Chinook salmon Chinook salmon (1999-2009) N = 20 31 26 19 N = 39 101 8 100 100

80 Euphausiids 80 Euphausiids Crustaceans Crustaceans Squids Squids Anchovy Anchovy 60 Smelt 60 Smelt Rockfish Rockfish Herring Herring Hake Hake 40 Other Fishes 40 Other Fishes

Percent weight contribution weight Percent 20 20

0 0 1981 1982 1983 1984 May June September

(Brodeur and Pearcy 1992) (Daly and Brodeur unpub.) Available Forage Fish Prey Sizes from Trawl Surveys (1998-2011)

Whitebait Pacific Smelt Sardine

Northern Pacific Anchovy Herring

Represents > 90% of all forage fish collected in pelagic trawl surveys done off the Columbia River at night

Brodeur, Buchanan, and Emmett (2014) CalCOFI Rep. Overall Forage Fish Prey Sizes from Trawl Surveys (1998-2011) (Scaled to Relative Abundance)

All Forage Fish Lengths Combined

Represents > 90% of all forage fish collected in pelagic trawl surveys done off the Columbia River at night Brodeur, Buchanan, and Emmett (2014) CalCOFI Rep. Fish Prey Sizes eaten by Pacific Hake and Chinook Salmon

140 100 Pacific hake (1998-2011) Chinook salmon adults (1998-2004) 120 Anchovy 80 Anchovy Herring Herring 100 Rockfish Rockfish Smelt Smelt 60 80 Sardine Sardine Other Other 60

Frequency Frequency 40

40 20 20

0 0 0 50 100 150 200 250 0 50 100 150 200 Prey size class (mm) Prey size class (mm)

Brodeur, Buchanan, and Emmett (2014) CalCOFI Rep. Ranking of Top Three Predators on Each Forage Fish

Pelagic Fishes Demersal Fishes Elasmobranchs Based on proportion of diet made up by forage species Anchovy Sardines Herring Smelt unid. Hake juv. Rockfish juv. 1 Albacore Chinook Boccacio Black rockfish Arrowtooth Sablefish 2 Coho Spiny dogfish Spiny dogfish Chinook Soupfin shark Yelloweye 3 Jack mackerel Soupfin shark Chinook Yelloweye Halibut Halibut

Proportion of diet scaled to total biomass in NCC Anchovy Sardines Herring Smelt unid. Hake juv. Rockfish juv. 1 Albacore Hake Hake Hake Hake Sablefish 2 Jack mackerel Spiny dogfish Spiny dogfish Black rockfish Spiny dogfish Hake 3 Spiny dogfish Albacore Jack mackerel Lingcod Albacore Jack mackerel

Brodeur, Buchanan, and Emmett (2014) CalCOFI Rep. Data Required to Estimate Salmon Forage Fish Consumption

• Salmon diet analysis: annual time series of prey types consumed from BPA seasonal surveys for Chinook and Coho, 2004-2011. • Salmon abundance: annual time series of abundance for 3 months from BPA seasonal Salmon Surveys, 2004-2011. • Prey abundance: annual time series of late larval/juvenile rockfish and anchovies abundance per tow from NMFS Juvenile Surveys, 2004-2011. • Individual consumption estimates: annual (2004-2011) time series of bioenergetic model runs using prey abundance, growth rates, and temperature in the upper 20 m integrated across the shelf.

5 Top-down effects by salmon on focal prey

Juvenile Rockfish Consumption 60

50 June Chinook Salmon Coho Salmon 40 Mean _ 30 X = 10.9 20 _ 10 X = 15.8

Percent Percent Consumed 0 2004 2005 2006 2007 2008 2009 2010 2011 60 Years 50 September _ 40 X = 12.9 30 _ 20 X = 2.1 10

Percent Percent Consumed 0 2004 2005 2006 2007 2008 2009 2010 2011 Years

Brodeur and Daly (unpub.) Juvenile Anchovy Consumption 20 June 15 Chinook Salmon Coho Salmon 10 Mean _ 5 X = 4.9 _ 0 X = 1.8

Percent Consumed Percent 2005 2006 2007 2008 2009 2010 2011 30 September Years

20 _ X =12.4 10 _ X = 1.1 0

Percent Consumed Percent 2005 2006 2007 2008 2009 2010 2011 Years

Brodeur and Daly (unpub.) Data Gaps • Diet information available only through ancillary collections so big data gaps in space and time (i.e., few dedicated process surveys to look at predator- prey interactions) • Little sampling outside of summer season and in nearshore ecosystem • No information on prey selectivity by predators and how that changes with prey availability • Need more information on prey quality in terms of spatial and temporal variability

U.S. Department of Commerce | National Oceanic and Atmospheric Administration | National Marine Fisheries Service | Page 32 Lipid Densities Vary Between Months and Years

8 30 May AnchovyMonthly Lipid Values (% June Herring 25 WhitebaitWt. Weight) 6 Sardine

20

4 15

% Lipid % Total% Lipids 10 2

5

0 0 JuneES05 05 AugLS05 05 JuneES06 06 AugLS06 05 Time Period Gadid WARM AVERAGE Cancer Ronquil Hyperiid Osmerid Rockfish Euphausiid Hexagrammid

Marine Biology (2010) 157:1975–1987 Fatty acid profiles of juvenile salmon indicate prey selection strategies in coastal marine waters Elizabeth A. Daly · Cassandra E. Benkwitt · Richard D. Brodeur · Marisa N. C. Litz ·Louise A. Copeman

33 Acknowledgements • The crew and scientists of the NOAA Ship Bell M. Shimada, R/V Ocean Starr and F/V Frosti • NOAA and Bonneville Power Administration for funding