Quesnel Lake Sockeye Salmon:

Quesnel Lake Sockeye Salmon: Large escapements, trophic surprises, and the consequences of density-dependence Daniel T Selbie, Ph.D. Cultus Lake Salmon Research Laboratory Science Branch, Fisheries and Oceans Canada Quesnel Lake, BC Cariboo Region

Fraser River Sockeye Salmon Returns Hell s Gate Slide Total Fraser River Returns 1893-2011 Cohen Inquiry Report I, 2012

Declining Fraser River Sockeye Production Cohen Inquiry Report I, 2012

Freshwater Environmental Limitation of Stock Productivity Density-Dependent Factors Spawning Limitation Rearing Limitation Density-Independent Factors Disease Inter-Specific Competition Predation Environmental Variability Environmental Stressors Adams River 2010, Photo: R. Bailey, DFO

Major stocks have exhibited spawning escapements in excess of modelled optimal escapements Selbie et al. 2010, PSC Workshop Proceedings

Rearing Limitation Sockeye Planktivory Nutrient Stimulation

Marine-Derived Nutrients

Phosphorus Bottom-Up: Marine-Derived Nutrients in Lakes Nursery Lake Fertilization Lake Dalnee, Russia Lake Blizhnee, Russia Salmon Escapement Krokhin 1975. Ecol. Studies. Hyatt et al. 2004. Environ. Rev. 12: 133-162

Juvenile Productivity Index Density Dependence Shuswap, Quesnel, Chilko lakes Evidence of densitydependent survival across escapement densities in Shuswap, Quesnel and Chilko lakes Fall Fry Abundance (Millions) Fall Fry Abundance (Millions) 200 150 100 50 0 80 60 40 20 0 Shuswap Fall Fry vs. EFS 0.0 1.0 2.0 3.0 Quesnel r 2 =0.38* Fall Fry vs. EFS r 2 =0.33* 0.0 1.0 2.0 * Data include dominant and subdominant cycle years but not non-dominant years Smolt Abundance (Millions) 80 60 40 20 0 Chilko 2005 2006 Smolts vs. EFS r 2 =0.36* 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 EFS (Millions) Selbie et al. 2010, PSC Workshop Proceedings

Zooplankton Reality: It s Top-Down & Bottom-Up Global Dataset: r 2 = 0.33 *, p < 0.001, n = 82 Inter-Lake: r 2 = 0.46 *, p = 0.03, n = 11 Sockeye Planktivory Nutrient Stimulation Salmon Chen et al. 2011. Oikos 120: 1317-1326.

Quesnel Lake, BC Selbie and Shortreed, in prep.

North Arm Main Arm East Arm Quesnel Lake Selbie and Shortreed, in preparation

North Arm Main Arm East Arm Quesnel Lake Selbie and Shortreed, in preparation

Lake Responses to Variable Escapements Water chemistry and primary productivity responses Selbie and Shortreed, in preparation

Marine-Derived Nutrients Stable Nitrogen Isotope (δ 15 N) Tracer Zooplankton North Arm Main Arm East Arm Quesnel Lake Juvenile Sockeye Selbie and Shortreed, in preparation

Nursery Lake Primary Productivity Seasonal Mean Photosynthetic Rates Lake productivity appears to have increased in Quesnel Lake since 1980 s & 1990 s Selbie et al. 2010, PSC Workshop Proceedings

Juvenile Sockeye Salmon In-Lake Growth Juvenile Fall Fry Weight * 27-45% fry weight reduction in 2001-02 BY Juvenile Fall Fry Length * 7-16% fry length reduction in 2001-02 BY

Freshwater Survival Index Fall Fry/Effective Female Spawner S D S D D D S D S D S D S D Dominant Cycle Line S Non-Dominant Cycle Line

Autotrophic Phytoplankton & Bacteriaplankton Composition & Production by Functional Size Class Selbie and Shortreed, in preparation

Diatoms: Tabellaria spp. A trophic dead end? T. fenestrata Selbie and Shortreed, in preparation

Rhizosolenia eriensis Lake P-fertilization McQueen et al. 2007. N. Am. J. Fish. Mgmt. 27: 369-386.

Zooplankton Biomass Fall Total Zooplankton and Daphnia North Arm Main Arm East Arm Quesnel Lake Selbie & Shortreed, in preparation

Zooplankton Size Variation Dominant taxa North Arm Main Arm Quesnel Lake East Arm Selbie and Shortreed, in preparation

Juvenile Sockeye Diet Stomach content frequency and biomass Proportion by frequency (%) Proportion by weight (%) 100 80 60 40 20 0 A) Fall fry diet by frequency of occurrence 1994 1995 1999 2000 2002 2003 2004 Brood Year 100 80 60 40 20 B) Fall fry diet by biomass 0 1994 1995 1999 2000 2002 2003 2004 (S) (N) (N) (N) (S) (N) (N) Daphnia Bosmina Eubosmina Epischura Leptodiaptomus Diacyclops Insect Other (N) Non-Dominant Cycle Line (S) Sub-Dominant Cycle Line

Juvenile In-Lake Growth Juvenile Fall Fry Weight * 27-45% fry weight reduction in 2001-02 BY Juvenile Fall Fry Length * 7-16% fry length reduction in 2001-02 BY

Size Matters: Smolt to Adult Survival Dependency upon freshwater growth Koenings et al. 1993. Can. J. Fish. Aquat. Sci. 50: 600-611

Log Returns/1000 Fry Log Returns/1000 Fry Inter-Stage Freshwater Growth Effects Post-Fall Sockeye Survival vs. Length & Weight 2.5 2 r 2 = 0.45, p = 0.008 r 2 = 0.54, p = 0.003 2.5 2 1.5 1.5 1 1 0.5 Weight 0.5 Length 0 0 1 2 3 4 5 6 Fall Fry Mean Weight (g) 0 50 60 70 80 Fall Fry Mean Length (mm)

Overwinter & Marine Survival to Adult Adult Returns/Fall Sockeye Fry S S D S D D D D S S D

Delayed Density Dependence (DDD) Delayed Density-Dependence (DDD) (Brood Year Interactions) Large escapements in a brood year negatively impact the brood and at least the following three broods (Peterman & Dorner 2011) Explicit in Larkin S/R Model A proposed explanation for cyclic dominance in Fraser sockeye Hypothetical Mechanisms for DDD Dominant Cycle Line (1) Simple density-dependent mechanisms (dominant brood year) Successive Cycle Lines (2-4) Disease on densely populated spawning grounds Increased reproduction and survival of long-lived sockeye predators Severe inter-annual depletion of nursery lake food webs

DDD: Fraser Stock-Recruit Evidence Larkin vs. Ricker (Peterman & Dorner 2011) Ricker Model Stationary stock recruit Larkin Model Ricker model with cycle line interaction Larkin better fit in 9 of 19 stocks Quesnel, Scotch, Stellako Larkin model best fit Conclusion - DDD occurs in some stocks, but not all; Quesnel most pronounced evidence Modified from Peterman and Dorner 2011, Cohen Commission Report #10

Evidence of Cycle Line Interactions Quesnel Lake Fall biomass (kg/efs). 2.0 Quesnel Lake Dominant Subdominant 1.0 y = -0.52Ln(x) + 7.44 R 2 = 0.91 y = -0.16Ln(x) + 2.27 R 2 = 0.53 0.0 10,000 100,000 1,000,000 10,000,000 Effective Female Spawners Horsefly River - Quesnel Lake Horsefly River Circuli count (C1) 15 2ndNonD - r 2 = 0.78 Dom - r 2 = 0.74 1stNonD - r 2 = 0.63 SubD - r 2 = 0.97 10 1,000 10,000 100,000 1,000,000 10,000,000 Effective Female Spawners Woodey, Lapointe & Hume. in prep

Are there lessons to be learned from Alaska?

Lessons from Alaska Large Escapements & Productivity (Clark et al. 2007) Long-term productivity declines & increased stock variability when escapement goals exceeded Believed to be linked to surpassing nursery ecosystem productive capacity Delayed Density Dependence (Clark et al. 2007) Detected DDD in 5 stocks with over-escapement R/S fell below replacement for 2-5 yr following consecutive over-escapements (> 2x S max )

Lessons from Alaska Barren Lakes Experiments (Koening & Kyle) Experimental sockeye introductions & fertilization Persistent Top-Down Effects Restructuring of zooplankton community by sockeye can result in a resilient & resistant food web reinforced by modest planktivory Severe/prolonged sockeye foraging can cause brood interaction & depression Fertilization can re-establish bottom-up control, but time lags evident and potentially very long

Lessons from Alaska Frazer Lake, AK Sockeye introduced due to fish latter Over-escapement resulted in collapse of dominant brood line and reinforcement of cyclic dominance Large escapements not independent (as per S/R assumptions) Freshwater food webs are likely the linkage

Quesnel Lake Fall Fry Ricker Ricker Rmax Smax PR Pred Record Escapements Fall fry (millions) 80 60 40 20 0 0.0 0.5 1.0 1.5 2.0 Effective female spawners (millions) Quesnel (2001-02) S/R adult : 187-223% S max S/R juvenile : 155-222% S max PR model: 280-334% S max Shuswap Lake Fall Fry Ricker Ricker Rmax Smax PR pred BY 2010 Fall fry (millions) 200 150 100 50 0 0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 Effective female spawners (millions) Shuswap (2010) S/R adult : 215% S max S/R juvenile : 366% S max PR model: 481% S max Smolts (millions) 80 60 40 20 0 Chilko Lake Smolts Ricker Ricker Rmax Smax PR pred BY 2010 Recent yrs PR <1996 PR 2009 0.0 0.2 0.4 0.6 0.8 1.0 1.2 Effective female spawners (millions) Chilko (2010-11) S/R adult : 204-547% S max S/R juvenile : 198-252% S max PR model: 151-357% S max

Quesnel Summary Apparent Changes in Carrying Capacity Forced by MDN variation at high escapements Apparent long-term increase in photosynthetic rates and PR model carrying capacity estimates Trophic Interactions, Freshwater Growth & Survival Trophic energy transfers can be largely interrupted by dead ends Top-down forcings erode bottom-up influences at high densities Ultimate impacts on juvenile condition and possibly latelake/marine survival, which may be persistent Delayed Density Dependence (DDD) Potential Peterman et al. (2010) & Peterman and Dorner (2011) found only Quesnel showed striking evidence to date Supporting evidence from juvenile data (Woodey et al. in prep) Are there concerns for other stocks?

Sockeye Salmon Conservation Units Lake-type CU River-type CU ~216 lake-type sockeye conservation units in Canada Holtby and Ciruna 2007, CSAS Res. Doc. 2007/070