Marine Survival of Puget Sound Chinook salmon- New studies on size-selective mortality and critical growth periods

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Western Washington University Western CEDAR Salish Sea Ecosystem Conference 2014 Salish Sea Ecosystem Conference (Seattle) May 1st, 1:30 PM - 3:00 PM Marine Survival of Puget Sound Chinook salmon- New studies on size-selective mortality and critical growth periods David Beauchamp University of Washington, davebea@uw.edu Follow this and additional works at: http://cedar.wwu.edu/ssec Part of the Terrestrial and Aquatic Ecology Commons Beauchamp, David, "Marine Survival of Puget Sound Chinook salmon-new studies on size-selective mortality and critical growth periods" (2014). Salish Sea Ecosystem Conference. 197. http://cedar.wwu.edu/ssec/2014ssec/day2/197 This is brought to you for free and open access by the Conferences and Events at Western CEDAR. It has been accepted for inclusion in Salish Sea Ecosystem Conference by an authorized administrator of Western CEDAR. For more information, please contact westerncedar@wwu.edu.

Marine Survival of Salmon: Size-selective Mortality and Critical Periods Dave Beauchamp University of Washington WA Cooperative Fish & Wildlife Research Unit Washington Sea Grant Pacific Salmon Commission Salmon Recovery Fund Board Nisqually, Tulalip, Skagit Coop. & Lummi Tribes NOAA, WDFW, Kwiaht, DFO-Canada, Pac. Salmon Foundation

Study Objectives Use Size-selective mortality to identify critical periods of growth that influence survival to adulthood Which life stages & associated habitats? FW, estuarine delta, marine nearshore, offshore? Diagnose what factors limit growth or survival during critical periods Food supply or quality Temperature Competition Predation Approach & example of pilot results

Survival Linked to Size & Growth at Specific Life Stages Size at release & Marine entry NOT Correlated to Surv. Marine survival Strongly linked to Wt after 1 month Epi-pelagic feeding In Puget Sound through July 2-3 fold Wt gain during 1 o pelagic feeding Weaker pattern In Sept. % Smolt-to-Adult Returns (SARs) 2.0 1.5 1.0 0.5 0.0 Duffy & Beauchamp 2011 CJFAS 68:232-240 Puget Sound age-0 CWT Hatchery Chinook Hatchery Release Wt r 2 = 0.09 2x Wt July Wt r 2 = 0.84 4x Surv. 0 10 20 30 40 50 60 Body mass (g) Critical growth period June-July offshore Hatchery Chinook Nearshore? Sept Wt r 2 = 0.55 Do critical periods vary among stocks? (Nisqually, Snohomish, Skagit, Nooksack) Critical periods in FW, estuarine, nearshore or offshore life stages?

Hatchery: pre-release size structure & scales release date & abundance Outmigrant Trap Timing, Abundance Size, Scales, (~Diet & Otoliths from morts) Weekly Feb/Mar to ~July Estuarine Channels (trap or B Seine) & Nearshore Beach Seine Timing, Abundance Size, Scales, Otoliths, Diet 2x per month Offshore Purse seining Timing, Abundance Size, Scales, Otoliths, Diet ~2x per month Including predatory fish mid-apr to mid-oct Offshore Midwater Trawl Depth-stratified 15-m depths Timing, Abundance Size, Scales, Diet Including predatory fish July & Sept Returning Adults: Scales & Otoliths & Residents

Track Specific Populations thru Early marine life stages Methodical near/offshore sampling For selected watersheds Size structure Timing & duration Total/relative abundance Diet & Body Condition Environmental Conditions Freshwater: Hatchery Release Smolt traps Estuarine & nearshore Marine: Estuarine Tide channel traps 2x/mo Beach seining 2x/mo Mar-Sep/Oct Offshore: Zooplankton 2x/mo Apr-Oct Purse seining 1-2x/mo May-Sep DFO Midwater Trawl: Sep Grants approved, no funding, DFO

Timing of Life Stage Pulses: Downriver Migrants Relative Abundance 1.0 0.8 0.6 0.4 0.2 River Migrants 0.0 Mar Apr May Jun Jul Aug Sep Oct Date

Timing of Life Stage Pulses: Estuarine Delta & Nearshore Relative Abundance 1.0 0.8 0.6 0.4 0.2 River Migrants Estuarine Delta & Nearshore 0.0 Mar Apr May Jun Jul Aug Sep Oct

Timing of Life Stage Pulses: Offshore (within Puget Sound) Relative Abundance 1.0 0.8 0.6 0.4 0.2 River Migrants Estuarine Delta & Nearshore Offshore w/in Puget Sound 0.0 Mar Apr May Jun Jul Aug Sep Oct Date

Relative Abundance 1.0 0.8 0.6 0.4 0.2 Size-Selective Mortality by Life Stage River Migrants Estuarine Delta & Nearshore Offshore w/in Puget Sound 0.0 Mar Apr May Jun Jul Aug Sep Oct Date Fork Length (mm) 200 150 100 50 Avg. Juvenile in field samples Feb Apr Jun Aug Oct

SSM: Scales used to back-calculate growth history of known-origin Juveniles & Adults scale radius ~ fish length circulus spacing ~ growth rate J. Boldt Age-0 Chinook BY 2001, June 2002 in PS Circuli from Same Growth Region for Juv. & Adult from same brood year

Relative Abundance 1.0 0.8 0.6 0.4 0.2 Size-Selective Mortality by Life Stage River Migrants Estuarine Delta & Nearshore Offshore w/in Puget Sound 0.0 Mar Apr May Jun Jul Aug Sep Oct Date Fork Length (mm) 200 150 100 50 Surviving adults Size or Growth diverge during: Offshore growth Delta-Nearshore River growth Avg. Avg. Juvenile Juvenile in field samples in field samples Feb Mar Apr Apr May Jun Jul Aug Aug Sep Oct Oct

Size Selective Mortality (SSM) For each life stage: SSM Inferred by comparing Size distributions (scale radius at specific circuli) for juveniles to: -Juveniles at later life stages -Adult survivors from that same group Larger individuals survived at a Disproportionately higher rate. In this hypothetical example: -The Smaller 50% of juveniles contribute only 5% of the surviving adults If significant SSM observed between periods, Then diagnose factors affecting growth within these periods based on bioenergetics modeling Frequency 0.5 0.4 0.3 0.2 0.1 0.0 5% 0 2 4 6 8 10 0.5 Scale Scale radius radius units at B Circuli # 0.4 A Size-selective mortality indicated by shift in size at a specific circulus # All Juveniles Survivors: No selection All Juveniles at 0.3circuli #s corresponding to: Alternate Life history? -Hatchery 0.2 release Survivors -Delta 0.1 & Nearshore Marine rearing Survivors SSM 0 2 4 6 8 10 12 Scale radius at a specified circulus SSM Examine scale-based size distributions -Smolt outmigration -Offshore rearing -Returning 0.0 Adults 50%

Higher Feeding Rate = Higher Growth & Survival Body Mass (g) 30 20 10 97 mm North; S = 0.7% Central; S = 0.6% South; S = 1.0% High Survival (0.8%) 3-4 x Wt increase offshore after peak nearshore use Release Hatchery PS Chinook (CWT groups) 2001 143 mm 134 mm 124 mm 96 mm North; S = 0.5% Central; S = 0.5% South; S = 0.3% Low Survival (0.4%) 2x Wt increase offshore after peak nearshore use 2002 120 mm 117 mm 0 86 mm 84 mm Nearshore Offshore 5/7 5/21 6/4 6/18 7/2 7/16 86 mm 91 mm Nearshore Offshore 5/7 5/21 6/4 6/18 7/2 7/16 Consumption (g/period) 150 100 50 20% higher avg feeding rate S=0.6-1.0% 66-90% C max 64% C max g.e. = 16% 90% C max g.e. = 16% Insects 88% C max g.e. = 14% Crab Larvae 2001 Copepod Polychaete 2002 Gammarid Fish Hyperiid Crab larvae Insect Euphausiid Other Invert S=0.3-0.5% 55-64% C max Low & Variable Feeding rates Suggest food limitation-56% C max 62% C max 65% C max g.e. = 12% Prerequisite g.e. = 16% g.e. for = 15% competition 0 Duffy 2009 Dissertation North Central South North Central South

Temperature Effects on Metabolism, Feeding & Growth Daily Growth Rate (g g -1 d -1 ) 0.030 0.025 0.020 0.015 0.010 0.005 0.000-0.005 10-g Chinook salmon 2800 J/g diet 0 5 10 15 20 Beauchamp & Duffy 2011. PSC Rept Spr-Sum Epi-pelagic Temps in PS Spr Sum Temperature o C 100% C max 75% C max 50% C max 25% C max SoG Upper Temp Temperature has Less effect on Summer Growth than feeding rate for PS Chinook Thermal effects are Likely more important In Strait of Georgia Thermal Tolerance

Summary Approach Identify Critical Periods: Let the fish tell us what s happening! Critical periods can vary among Spp & Stocks Methodical Sampling: Hatchery & smolt traps in FW, estuarine & nearshore marine, local epi-pelagic, open ocean, Adult returns Use SSM to identify critical periods and associated habitats (lengths, scales &/or otoliths) Diagnose factors affecting growth & survival within critical periods Can Inform restoration efforts & priorities

Size-Selective Mortality Size-selective Mortality (SSM) is Prevalent Stage-specific size positively correlated to survival Can be used to identify critical periods of mortality or growth which influences mort SSM differs among Spp, stocks, life stages Shifts in size-at-age distribution among life stages reflects timing & magnitude of SSM Need to account for stock origin & migration SSM doesn t rely on serial abundance est. SSM can link top-down & bottom-up factors affecting Survival & Growth

Marine Survival trends for Hatchery Chinook Marine survival of hatchery Chinook in the Salish Sea region declined 1980s & remained low-but not adjacent regions 2 CHINOOK Strait of Georgia Puget Sound 1.5 Marine survival ( % ) 1 0.5 0 1982 1985 1988 1991 1994 1997 2000 2003 Year to Sea Beamish. 2011.

BACKGROUND: Puget Sound Hatchery Chinook Salmon Marine Survival Rates (smolt-adult) by release year # CWT groups examined 1.5 2 2 2 7 9 8 62 7 8 8 7 8 8 6 8 3 6 4 10 10 4 4 4 CHINOOK Strait of Georgia Puget Sound 1.5 Marine survival ( % ) 1 0.5 0 1982 1985 1988 1991 1994 1997 2000 2003 Survival (%) 1.0 Year to Sea 0.5 0.2-1.0% 21/23 yrs 0.0 1984 1986 Ruggerone & Goetz 2004 Duffy This 2009 study 1988 1990 1992 1994 1996 Release Year 1998 2000 2002 2004 2006

Scale radius (microns) Incremental circuli spacing (microns) Back-calculated size-at-age of juvenile & adult Puget Sound Chinook 600 500 400 300 200 60 55 50 45 40 35 30 Demonstration of Concept: Size of Juv Chinook Offshore approximates Size & G for Surviving Adults 3 6 9 12 15 18 21 Nearshore Offshore Adults Circulus number Growth rate of Juv Chinook Offshore approximates that for Surviving Adults. ± 2SE Juv. nearshore 2002 Juv. Offshore 2008 Adults 2004 Juveniles offshore Juveniles nearshore May-Jun (10-12) July (16-18) Sept (19-21) Growth period (Circulus interval) ± 2SE SSM & Critical period approach: Compare size-specific contributions of different life stages to adult returns & diagnose limits to growth & S in Critical periods Questions for 2014-2015: -Timing of life stages pulsing through specific habitats -Size-specific contribution to later stages -Identify critical life stages & assoc.habitat -Growth performance w/in life stage/habitat -Diagnose factors limiting growth: -Food supply (production, competition) -Food quality (energy content) -Thermal effects on metabolism -SSM Predation impacts by resident salmon

Size Selective Mortality (SSM) For each life stage: SSM Inferred by comparing Size distributions (scale radius at specific circuli) for juveniles to: -Juveniles at later life stages -Adult survivors from that same group Larger individuals survived at a Disproportionately higher rate in Both hypothetical examples A & B B, 2 nd mode of smaller juveniles did not survive to Adulthood. This mode could represent growth trajectories from an alternative life history strategy that s unsuccessful under prevailing conditions Frequency 0.5 0.4 0.3 0.2 0.1 0.0 0.5 0.4 0.3 0.2 0.1 0.0 A Size-selective mortality indicated by shift in size at a specific circulus # All Juveniles Survivors: No selection 0 2 4 6 8 10 B Scale radius units All Juveniles Alternate Life history? Survivors Survivors SSM 0 2 4 6 8 10 12 Scale radius at a specified circulus SSM

Rapid Early Marine Growth Improves Survival 1.5 1.2 Similar relationship for all offshore juvenile Chinook during July (but ~80% hatchery) Survival (%) 0.9 0.6 1997-2005: 4-fold difference in ocean survival '99 '04 '97 '01 '05 '00 0.3 '02 '98 0.0 2x difference in July Wt 10 15 20 25 30 35 July Wet Weight (g)

Potential Pelagic Competitors During Critical Period: Pacific Herring dominate the biomass of epi-pelagic planktivores Biomass & Spatial Temporal Overlap July 2004 15 Daylight Planktivore Community Mean Catch/hr 0 5000 10000 15000 20000 25000 30000 35000 Shallow: 0-15 m Herring: smaller aggregations at greater depth (scattering layer) Depth (m) 30 45 60 Chinook Other Salmon Herring Other DFO Canada Midwater trawl

Length frequency Temporal Diet Composition Thermal Experience 120 100 80 60 40 Modeling Process Consumer Growth Bioenergetics Model Predator Energy Density Prey Energy Density 20 0 200 300 400 500 600 Fork length (mm) Consumer Size Structure & Abundance Consumption Estimate Population Consumption by Herring & Chinook Compare Consumption Demand by Herring & Chinook for Key Prey During Critical Growth Period (May-July)

Potential Inter-specific Competition for Food Herring remove 10-47x more Biomass of key shared prey than H+W Chinook during Critical May-July period CONCLUSION: On average, Competition driven 1 o by Herring in pelagic Habitats of Puget Sound. Prey consumption demand (MT/season) 20000 18000 16000 14000 12000 10000 But-Competition should be Considered across the entire Epi-pelagic planktivore community 8000 6000 4000 2000 0 37x Copepod Crab Larvae Population-level consumption demand 2008: SPS, CPS, NPS 10x Intensity of competition will likely Vary among regions & months, based on relative abundance & diet of each species 45x Euphausiid Gammarid by Herring age 0-4 by Chinook age-0 47x Hyperiid Other