Adult Sockeye Migrations: Challenges to Complete the Journey and Spawn Scott Hinch University of British Columbia Vancouver, BC, CANADA Outline of Presentation 1. Ocean conditions matter for freshwater migrations 2. Populations are unique 3. Entry timing and physiological condition 4. Warming temperatures and climate change • Fraser sockeye starting upriver migration are heavier when ocean food more abundant Hinch et al 1995 • Fraser sockeye starting upriver migration have higher energy reserves when ocean temperatures are cool and upwelling is strong Crossin et al 2004 • spawner to recruit survival is lower when coastal temperature is warmer Hinch et al 1995 Outline of Presentation 1. Ocean conditions matter for freshwater migrations 2. Populations are unique 3. Entry timing and physiological condition 4. Warming temperatures and climate change Fraser River Sockeye Salmon ~ 150 stocks which differ in: • spawning migration distance (100 – 1,200 km) • annual spawning abundance (100’s – 1,000,000’s) • adult morphometry, energy density, fecundity, swimming performance, heart morphology Adaptations for completing river migration and spawning Adults with long distance or high elevation spawning areas have: -small torpedo shaped bodies -high body energy -few and small eggs -high maximum swimming speeds -more efficient hearts -large and broad ‘metabolic scope’ ‘Difficult Migrations’ -built for high performance and energy conservation ‘Easy Migrations’ Swimming performance and metabolic limits • Swim tunnel respirometry studies Metabolic scope & temperature Maximum scope Oxygen consumption (Mo2) Maximum Mo2 Zero scope Metabolic scope Topt Routine Mo2 (Q10 effect) Tcrit Temperature Metabolic scope = O2 available for activities other than routine and is temperature-dependant Metabolic scope (mg O2 kg-1 min-1) Metabolic scope temperature profiles for 3 Fraser salmon stocks 12 A 10 8 Gates Creek Sockeye 6 4 Topt Gates Creek 2 (summer-run sockeye) 0 B Weaver Creek 10 (late-run sockeye) 8 Weaver Creek Sockeye Chehalis River 6 (fall-run coho) Tcrit 4 Topt 2 0 Key points C 10 • Topt and Tcrit vary among stocks 8 Chehalis Coho 6 • Only 6-7oC between Topt and Tcrit 4 • Maximum scope varies among stocks in 2 Topt relation to migration difficulty 0 0 5 10 15 20 25 Lee et al. 2003 Temperature ( oC) Historical Lower Fraser River thermal conditions E. Stuart Early Summer Summer Temperature (°C) 22 Late Summer 20 18 1 SD 16 1940-2000 average 1 SD 14 12 26 3 10 17 24 31 7 14 21 28 4 11 Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Date temperature frequency distribution Metabolic scope (mg O2 kg-1 min-1) Metabolic scope temperature profiles for 3 Fraser salmon stocks 12 16% A 10 12% 8 Gates Creek Sockeye 6 8% 4 4% 2 Gates Creek (summer-run sockeye) 0 0% B 10 Weaver Creek 12% (late-run sockeye) 8 Weaver Creek Sockeye 6 8% Chehalis River (fall-run coho) 4 4% 2 0 0% C 10 12% Key point 8 Chehalis Coho • Topt is coincident with 6 8% historic river temperature profile 4 4% during the migration period 2 0 0% 0 5 10 15 20 25 Farrell et al. 2008 ) Temperature (oC Outline of Presentation 1. Ocean conditions matter for freshwater migrations 2. Populations are unique 3. Entry timing and physiological condition 4. Warming temperatures and climate change Early Migration of Late run stocks • 1995 to present – segments of all late run stocks have migrated upriver 4-6 weeks early than usual • In recent years late runs have nearly completely eliminated their estuarine holding behaviour • correlated with early migration has been extremely high mortality in freshwater Weaver Sockeye Timing and Mortality 270,000 fish Migration Mortality (% total run) 100 80 60 165,000 fish 2001 2000 490,000 fish 1999 1998 2003 2006 2007 2004 1996 1997 1995 2005 2002 1995 - 2007 40 1974 - 1994 20 0 Up-River Migration Date (50% past hydroacoustic facility near river mouth) Inter-university / Inter-agency research began 2002 Questions: Why are they migrating early? Why are they dying at such high levels in-river? Relevance to other runs and species? Study methods: • destructive sampling of fish captured along ocean and freshwater migratory routes for physiological and genomics work (~4,000 fish sampled) • radio and acoustic telemetry with biopsy assessments on individual fish (~2,000 fish tracked) • swimming performance and simulated migration experiments Acoustic receiver Radio receiver and antennae Acoustic transmitter i-button temperature logger Radio transmitter Transmitters inserted down throat into stomach on migrating adults The image cannot be displayed. Your computer may not have enough memory to open the image, or the image may have been corrupted. Restart your computer, and then open the file again. If the red x still appears, you may have to delete the image and then insert it again. Mechanisms of mortality: searching for physiological answers with ‘biopsy’ telemetry • Gross somatic energy assessed by microwave energy meter • blood sample to examine range of plasma ions, reproductive and stress hormones • tissue removed from first gill arch ionoregulatory function and for functional genomics assessments • small muscle plug also taken for functional genomics Cooke et al. 2005. Cooke et al. 2008. Telemetry lines, stations and adult salmon tagging sites - Partnership with POST, Kintama, and LGL Ltd Biopsy telemetry results reveal that early migrants are: i) physiologically stressed ii) more reproductively mature iii) dysfunctional ion regulation Causes of early migration not yet resolved: • evidence supporting environmental, physiological and behavioural hypotheses • advanced maturation suggests changes to internal and/or external migratory cues and an ‘open ocean’ link (or earlier) • en-route mortality is associated with higher than usual river temperatures Cooke et al. 2006a,b Crossin et al. 2009 Hinch 2009 Outline of Presentation 1. Ocean conditions matter for freshwater migrations 2. Populations are unique 3. Entry timing and physiological condition 4. Warming temperatures and climate change Potential causes of thermal-based migration mortality • metabolic collapse Fast processes (hours-days) -critical temperature limits • stress • disease • energy depletion Slow processes (days-weeks) -accumulated degree-day limits Collapse of metabolic scope occurs in warm years 2004 temperatures during migration - red bars D 80% 60% 40% 20% 0% 100% E 80% 60% 40% 20% 0% 0 temperature frequency distribution 100% Percentage of maximum aerobic scope 25% 20% 15% 10% 5% 0% 25% 20% 15% 10% 5% 0% 25 30% Gates Weaver Temperature (oC) 10 15 20 Key findings • River temperature can exceed Tcrit for some sockeye stocks • in 2004, expect high mortality for segment of Weaver run that experienced temperatures ≥ 20 C • telemetry confirmed > 70% of total run perished that year Farrell et al. 2008 Mathes et al. 2010 Laboratory thermal holding experiments • collect migrating fish as they enter the Fraser and hold in lab for their migratory period under different temperatures • simulate migrations under different thermal and stress conditions Lab experiments with adult late run sockeye Mortality (%) 100 90 80 70 60 50 40 30 20 10 12oC 18oC Importance of thermal refuges? 18oC 20oC 0 12oC 8 oC 15-16oC 0 5 10 15 20 25 River migration duration Days 30 35 Stefan Larsson unpub. data Crossin et al. 2008 Migratory Disease Issues • parasites, bacteria and fungal infections are temperature dependent • bacterial infections – e.g. Columnaris disease (Flavobacterium columnare) • fungal infections – e.g. (Saprolegnia sp.) 2004 Columnaris infection in Weaver / Harrison Sockeye in 2008 Parasite infections - e.g. Parvicapsula minibicornis • kidney infection in lab held Weaver Creek sockeye increased rapidly when individuals had accumulated ~ 370 degree days Maximum score of 25 In kidney glomeruli Severity of infection 25 20 15 10 5 0 0 100 200 300 Degree days 400 500 600 Wagner et al. 2006 Mortality in relation to accumulated degree days • Weaver Creek sockeye captured at river entry (24 days at 18 ºC) • mortality begins ~ 300 degree days (spawning ground arrival) • 70% mortality at ~ 500 degree days 60 25 50 20 40 15 30 10 20 5 10 0 Severity of infection Cumulative mortality (%) 70 0 90 180 270 360 450 540 Accumulated degree days ºC Wagner et al. 2006 Crossin et al. 2008 How important are cool water refuges? Thermal experience of an Adams sockeye 2006 from its recovered temperature logger Tagged Spawning grounds Hourly Temperature C 20 17 14 BC USA 100 km 11 Key findings: - No thermal refuges in lower Fraser 8 Ocean 5 21-Aug 31-Aug Fraser R. 10-Sep Thompson R. Thompson R. Kamloops L. Shuswap L. 20-Sep 30-Sep 10-Oct Adams R. 20-Oct - Adams sockeye use lakes as thermal refuges Weaver Creek migration • directed swim up Fraser River (5-6 days in August/September) • 1-30 day migration / holding in Harrison River or Lake • spawning begins in early October Harrison Lake Weaver Creek Harrison River Vancouver N 10 km Fraser River Canada USA Hourly depth profiles for 4 Weaver sockeye ‘holding’ in Harrison Lake Fraser River migration thermal issues: 1. Since 1940s, ~ 2 ºC increase in peak summer water temperature 2. Recent years have had extreme (record) high temperatures 3. Since 1996, Late-run sockeye enter river 4-6 weeks earlier than normal and thus encounter temperatures ~ 5 C warmer 4. Climate models predict at least a 2 C increase in summer water temperatures 10 12 Temperature ( oC) 14 16 18 20 Since 1940s, ~ 2 ºC increase in peak summer water temperature 1950 1960 1970 1980 Years 1990 2000 Dave Patterson, DFO Env. Watch Program Historical Lower Fraser River thermal conditions E. Stuart Early Summer Summer Temperature (°C) 22 Late Summer 20 18 1 SD 16 1940-2000 average 1 SD 14 12 26 3 10 17 24 31 7 14 21 28 4 11 Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Date Recent years have had extreme (record) high temperatures E. Stuart Early Summer Summer Temperature (°C) 22 Late Summer 2004 20 18 16 60-y ave 14 12 26 3 10 17 24 31 7 14 21 28 4 11 Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Date Historical Lower Fraser River thermal conditions E. Stuart Early Summer Summer Temperature (°C) 22 Late Summer 2004 20 18 16 60-y ave 14 12 26 3 10 17 24 31 7 14 21 28 4 11 Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Date What about future conditions in the Lower Fraser River? E. Stuart Early Summer Summer Late Summer Temperature (°C) 22 2004 20 18 16 60-y ave 14 12 26 3 10 17 24 31 7 14 21 28 4 11 Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Date • 2004: 50-80% migration mortality across all runs • the future for Fraser River sockeye? E. Stuart Early Summer Summer Temperature (°C) 22 Late Summer 2004 20 Future 60-80 yr average 18 16 14 12 26 3 10 17 24 31 7 14 21 28 4 11 Jun Jul Jul Jul Jul Jul Aug Aug Aug Aug Sep Sep Date Model-averaged survival rate Survival rates (+/- CI) to natal rivers of fish with transmitters in relation to encountered Fraser River temperature (2002-2007) 1.0 1.0 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 0.1 0.0 0.1 Chilko 15 16 0.0 17 18 19 20 Quesnel 15 16 Model-averaged survival rate Temperature (oC) 1.0 0.9 0.9 0.8 0.8 0.7 0.7 0.6 0.6 0.5 0.5 0.4 0.4 0.3 0.3 0.2 0.2 0.0 Stellako-Late Stuart 15 16 17 18 19 Temperature (oC) 18 19 20 Temperature (oC) 1.0 0.1 17 20 0.1 0.0 Adams 13 14 15 16 17 18 19 Temperature (oC) 20 Martins et al. 2010 Predicting future migration survival with climate change models Temperature (oC) Summer-run Late-run (Adams) early-timed ~2 oC ~1 oC normal-timed normal-timed Quesnel Survival rate ~5 oC early-timed Stellako-Late Stuart ~1% ~16% ~15% Martins et al. 2010 Hague et al. 2010 Climate change summary: • warmer, less productive northeast Pacific ocean • reduction in preferred marine thermal habitat • poorer ocean survival • smaller mature fish with less energy • warmer river temperatures, more frequent ‘extreme’ years • higher rates of river migration mortality • effects will be stock-specific • thermal refuges will be key to survival of migrating sockeye in a ‘warming’ Fraser River particularly for stocks with lower Tcrit or disease thresholds Thanks to: • several past and current grad students, post-docs, technicians and colleagues • funders: NSERC, Pacific Salmon Commission, Fisheries and Oceans Canada All references can be found at: http://faculty.forestry.ubc.ca/hinch/ Research Team Ecology Physiology / Genomics Behaviour / Movement Parasitology Oceanography Scott Hinch (Univ. BC) Steve Cooke (Carleton Univ) Glenn Crossin (Univ. BC) Jeff Young (Univ. BC) Todd Mathes (Univ. BC) Tony Farrell (Univ. BC) Glenn Wagner (Univ. BC) Mark Shrimpton (Univ. Northern BC) Glen Van Der Kraak (Univ. Guelph) Kristi Miller (Can. Dept. Fish. Oceans) Michael Healey (Univ. BC) David Welch (Kintama and Pacific Ocean Shelf Tracking Project) Karl English (LGL Ltd.) Simon Jones (Can. Dept. Fish. Oceans) Stefan Larsson (Swedish Agricultural University) Rick Thomson (Can. Dept. Fish. Oceans) Fisheries Management Dave Patterson (Can. Dept. Fish. Oceans) Mike Lapointe (Pacific Salmon Commission) Take home messages: • Critical temperature limits (e.g. metabolic issues) and accumulated degree day limits (e.g. disease issues) can explain migration mortality • Provides mechanistic approach and understanding (predicting) potential effects of current and further climate change • Applying critical temperature or accumulated degree day limits will likely require stock- or run-specific information • Other relevant thermal related critical limits not presented: • Stress – e.g. plasma lactate > 18 mmol/L • Energy – e.g. somatic concentration < 4 MJ/kg • En-route thermal refuges are key to survival of migrating sockeye in a ‘warming’ Fraser River particularly for stocks with lower Tcrit or disease thresholds Biotelemetry of 2004 Weaver Creek sockeye Capture and tagging Harrison River Fraser River Tcrit Topt Todd Mathes M.Sc. thesis Key field findings - Critical temperature Issue • 0% survival of fish which experienced temperatures ≥ Tcrit • 10-30% survival of fish which experienced ‹ Tcrit › Topt • 50-80% survival for fish which experienced temperatures ~ Topt Capture and tagging dates Fraser River to spawning grounds Tcrit Topt Estimated degree-day accumulation for 83 Weaver sockeye from Fraser entry to spawning grounds based on telemetry and temperature reconstruction Degree-day accumulated (oC) a b b c Early-river Early-lake Normal-river Timing and Residency Groups Normal-lake Percent of time spent at depth for 6 adult sockeye in Harrison Lake Key Result – 80% of residency time spent at depths > 50 m and temperatures ~ 6-7 C Temperature manipulation field experiment • test hypothesis that high temperature-exposed fish (> 500 DD) survive poorly during migration compared to cool temperatureexposed fish (< 400 DD) • captured newly arrived, normal-timed, river migrants and held under different thermal regimes in captivity for three weeks, prior to release downstream of capture location Harrison Lake Weaver Ck. capture Harrison River release 10 km Fraser River Cold treatment (8 °C) 24 days 327 accumulated degree days N = 31 Warm treatment (18 °C) 24 days 510 accumulated degree days N = 17 • migration mortality higher in warm treated fish 70 • mortality rates reflect those of ‘early’ and ‘normal’ timed migrants Mortality (%) 60 50 510 degree days 40 30 327 degree days 20 10 0 78 80 Harrison Harrison Fraser Fraser Confluence Confluence 82 84 86 88 90 Distance from release site (km) 92 Spawning Spawning Grounds Grounds 94