Tom Reed (University College, Cork) Stocking and Climate Change

Salmon stocking and climate change
Tom Reed Beaufort Marine Research Award in Fish Population Genetics University College Cork, Ireland
Boosting salmon numbers - is stocking the answer or the problem? AST/IBIS conference
Glasgow November 2013
Talk outline
1. Physical changes 2. Signatures of climate change in salmon pop. dynamics 3. Reduced marine survival of Atlantic salmon 4. Freshwater impacts on Atlantic salmon 5. Case study of Burrishoole salmon (wild + ranched) 6. Resilience of populations and stock complexes in the
face of uncertainty
Global scale: atmospheric CO2 is increasing

Tripati et al. Science 2009
1. Physical changes
2. Signatures in salmon pops
3. Marine survival
4. Freshwater impacts
5. Burrishoole case study
6. Resilience
and the oceans are getting more acidic.
Source: http://www.nature.com/scitable/knowledge/library/ocean-acidification-25822734
"This [acidification] is unprecedented in the Earth's known history. We are entering an unknown territory of marine ecosystem change, and exposing organisms to intolerable evolutionary pressure. The next mass extinction may have already begun. From: IPSO State of the Ocean Report 2013: see http://www.stateoftheocean.org
1. Physical changes 2. Signatures in salmon pops 3. Marine survival 4. Freshwater impacts 5. Burrishoole case study 6. Resilience
Global scale: average temperatures rising at unprecedented rate
Source: Moritz and Agudo Science 2013

Source: http://news.stanford.edu/news/2013/august/climate-change-speed-080113.html
MOVE
1. Physical changes 2. Signatures in salmon pops
ADAPT
3. Marine survival 4. Freshwater impacts
OR DIE.
5. Burrishoole case study 6. Resilience
Source: http://news.stanford.edu/news/2013/august/climate-change-speed-080113.html
1. Physical changes
3. Marine survival
6. Resilience
Source: http://en.wikipedia.org/wiki/El_Nio
Temperature difference from baseline global mean
Pacific decadal oscillation
1. Physical changes
3. Marine survival
6. Resilience
Mantua et al. Bull Am Met Soc 1997

Climate change impacts on salmon

MARINE ENVIRONMENT FRESHWATER ENVIRONMENT
Death or stress due to high temperatures. Death or stress due to high temperatures. Increased energetic costs of migration Altered growth and maturation patterns Changes in food supply. Changes in predation. Increased disease risk and parasites.
ABIOTIC/DIRECT EFFECTS
Increased energetic costs of migration Altered growth and maturation patterns Changes in food supply.
BIOTIC/INDIRECT EFFECTS
Changes in predation. Increased disease risk and parasites.
Jonsson, B., and N. Jonsson. "A review of the likely effects of climate change on anadromous Atlantic salmon Salmo salar and brown trout Salmo trutta, with particular reference to water temperature and flow." Journal of Fish Biology 75.10 (2009): 2381-2447. Crozier, L. G., et al. "Potential responses to climate change in organisms with complex life histories: evolution and plasticity in Pacific salmon." Evolutionary Applications 1.2 (2008): 252-270. 1. Physical changes 2. Signatures in salmon pops 3. Marine survival 4. Freshwater impacts 5. Burrishoole case study 6. Resilience
Problems due to changes in ocean climate

Marine survival has decreased substantially in recent decades; especially affecting MSW fish but 1SW also affected, particularly from Southern European stock complex
2SW Non-maturing 1SW Maturing
Source: Friedland et al. 2013 In Press

Changes in marine productivity are associated with the Atlantic Multi-decadal Oscillation
Source: Friedland et al. 2013 In Press

What about freshwater impacts on regional/local scales?

The clear message from the Salmon Summit in this challenging global environment is to maximise the number of healthy wild salmon that go to sea from their home rivers, since management options in the ocean are limited.
there may be only limited opportunities to respond to further declines through management of the fisheries, as these have already been closed or greatly reduced.
Malcolm L. Windsor, Peter Hutchinson, Lars Petter Hansen and David G. Reddin. 2012. Atlantic salmon at sea: Findings from recent research and their implications for management. NASCO document CNL(12)60. Edinburgh, UK. 20pp.
4. Freshwater impacts 5. Burrishoole case study 6. Resilience
1. Physical changes
3. Marine survival
Managing salmon in the face of the uncertainty about future environmental changes will be challenging. The goal should be to protect the genetic diversity of the wild Atlantic salmon in order to maximise their potential to adapt to the changing environment. Consistent with a Precautionary Approach, where there are uncertainties there is a need for caution. The absolute priority should be to conserve the productive capacity of the resource.
Source: NASCO Salmon Summit report
So managing salmon in the face of CC will realistically be all about minimising impacts in freshwater, estuarine and coastal environments, where we have more direct control.
Should stocking be a part of the repertoire of management options, or do the risks outweigh the potential benefits?
1. Physical changes
3. Marine survival
6. Resilience
Resilience in the face of uncertainty

Resilience = sustained productivity despite major environmental change.
ECOSYSTEM PRODUCTIVITY STOCK COMPLEX or METAPOPULATION PRODUCTIVITY POPULATION PRODUCTIVITY

1. Physical changes 2. Signatures in salmon pops 3. Marine survival 4. Freshwater impacts
Rate of generation of biomass.
Recruits per spawner Abundance
6. Resilience
Environmental change
Perfect adaptation
Degree of local adaptation

(match between phenotype + environment)
Increasing maladaptation
Population abundance
(of naturally spawning fish)
Time
Environmental change
Stocking
Perfect adaptation
Degree of local adaptation

(match between phenotype + environment)
Increasing maladaptation
Population abundance
(of naturally spawning fish)
Time
Burrishoole Case study:

Introgression with captive bred fish
Longterm experiment (Burrishoole River)
The Model
Main factors determining egg
to smolt survival in Burrishoole survival)
% hatchery eggs in spawning cohort (Winter temperature for eggs & fry (Winter temperature for parr (-
survival)
survival) survival) survival)
Winter temperature for smolts (+
Interaction between % hatchery eggs and winter temperature (-
McGinnity et al. (2009). PRSB , 276:36013610
Observed changes in climate
Future projections
Resilience of stock complexes
ECOSYSTEM PRODUCTIVITY
STOCK COMPLEX or METAPOPULATION PRODUCTIVITY
POPULATION PRODUCTIVITY
1. Physical changes
3. Marine survival
6. Resilience
Sockeye Salmon habitat in Bristol Bay

9 major rivers
North Pacific Ocean
Bristol Bay
each with many populations
L.A. Rogers
Wood River
Salmon biological features are adapted to local
habitat conditions and how these filter climate
Lake beaches
Age diversity
1
2 3
Small streams
L.A. Rogers
Salmon landscapes are shifting mosaics of suitable habitat

(sensu Stanford et al. 2005)
Commercial fisheries for sockeye salmon in Bristol Bay have been sustained for over 120 years
50,000,000
Togiak
Number of sockeye salmon caught in Bristol Bay (1893-2010)
45,000,000 40,000,000
Ugashik Egegik Nushagak Naknek-Kvichak
catch Commercial catch Commercial
35,000,000 30,000,000 25,000,000 20,000,000 15,000,000 10,000,000 5,000,000 0
1893
1903
1913
1923
1933
1943
1953
1963
1973
1983
1993
data from ADFG
Year
2003
Variability in salmon increases the rate of fisheries closures

100 90 80 70 60 50 40 30 20 10 0
Bristol Bay (intact portfolio)
Fisheries are closed 4 years every century
Number of returning salmon (millions)
average minimum spawners
100 90 80 70 60 50 40 30 20 10 0
Bristol Bay (eroded portfolio)
Fisheries are closed 40 times every century
X X
X XX
X X
XX X X X XX X X X X X XX X
X XXX
Time (100 years)
Stability and productivity derive from diverse and changing habitat

Bristol Bay, Alaska
Pacific Northwest
To keep every cog and wheel is the first precaution of intelligent tinkering Aldo Leopold (Round River, 1953)
NOAA
Disease is the bullet that's killing the frogs, but climate change is pulling the trigger. Global warming is wreaking havoc on amphibians, and soon will cause staggering losses of biodiversity J. Alan Pounds 2006
Acknowledgments
IBIS/AST and the organisers Daniel Schindler (University of Washington, Seattle) for slides Phil Mc Ginnity for slides and great discussions always! Beaufort Marine Research Award in Fish Population Genetics
This Beaufort Marine Research Award is carried out under the Sea Change Strategy and the Strategy for Science Technology and Innovation (2006-2013), with the support of the Marine Institute, funded under the Marine Research Sub-Programme of the National Development Plan 20072013.
Extra slides
Are humans causing current climate change?

Quote from George Monbiots recent book:
If you reject this explanation [man-made climate change]
for planetary warming , you should ask yourself the following questions: 1. 2. Does the atmosphere contain carbon dioxide? Does atmospheric carbon dioxide raise the average global temperature? 3. Will this influence be enhanced by the addition of more CO2?
4.
Have human activities led to a net emission of CO2?
If you are able to answer no to any one of them, you should put yourself forward for a Nobel Prize. You will have turned science on its head.
Salmon returns to Bristol Bay are two times more reliable than the individual components of the portfolio
Bristol Bay
10
10
Age diversity
1
2 3
All the same age
10
Major rivers
10
Streams
10
Total return (relative)
0. 1
0. 1
0. 1
0. 1
0.01
0. 01
0. 01
0. 01
1
0. 01
1960
year
2010
Zooming in to more regional scales
The previous analysis was focussed at a very broad organizational scale (i.e. all salmon in the Atlantic, or N American, S European stock complexes, etc) Showed that there is this broad coherence in the responses of populations to large-scale climate drivers, but also showed differential responses of the S and N European stock complexes However, the synchronizing effects of climate can be reduced by heterogeneity in (a) the local expression of regional climate variation, (b) other extrinsic determinants of population dynamics, such as the density of predators or competitors, and (c) population traits and local adaptations which determine the sensitivity of populations to changes in their environment. While much focus has been placed on the synchronizing effects of climate, less attention has been paid to the possibility that populations may show sensitivity to different climatic drivers, even within the same geographic region or WATERSHED, due to genetic and phenotypic heterogeneity among populations and differences in the physical and biotic features of habitats that they occupy
1. Physical changes
3. Marine survival
6. Resilience
Scale and the detection of climatic influences on productivity
1. Physical changes
3. Marine survival
Rogers 4. and Schindler 2008 Global Change Biology Freshwater impacts 5. Burrishoole case study 6. Resilience
Climate variables affecting productivity of individual streams

Population unit
Bear creek Hansen creek Happy creek Ice creek Fenno creek Hidden creek Lynx creek
Nursery Lake
Alegnagik Alegnagik Alegnagik Alegnagik Nerka Nerka Nerka
Most important climate variable

Lake T second summer Lake T second summer SST second summer Lake T second summer SST second summer Lake T first summer Lake T first summer
Direction of effect
+ + + + +
Pick creek
1. Physical changes 2. Signatures in salmon pops
Nerka
3. Marine survival
+
6. Resilience
Weakened portfolio effect in a collapsed Chinook salmon population complex

Carlson & Satterthwaite 2011 CJAFS Sacramento R
San Joaquin R
One major question then becomes whether it is better to improve production in the SJ Basin by improving the environment, which may take a long time, or through hatchery production, which may foster homogeneity among rivers. Here we argue that restoring environmental heterogeneity, which is the template that gives rise to local adaptations and diverse life history portfolios, will pay larger dividends in the long run.
Climate variables affecting productivity at higher spatial scales

Population unit
Lake Alegnagik Lake Nerka All streams Wood river system
Number of lakes
1 1 2 5
Most important climate variable

Lake T second summer Lake T first summer Lake T second summer Date of ice breakup in year of smolting
Direction of effect
+ +
+
-
Inclusion of fall and winter climate variables was never strongly supported, at any scale Inclusion of the PDO (Pacific Decadal Oscillation) never well supported, even at level of entire Wood River system But evidence still for unexplained oscillations in productivity across years (over and above the identified climate effects)
Modelling effects of climate change: most appropriate level of spatial complexity to consider depends on specific management, scientific, or conservation goals
Inherent trade-off:
Coarse spatial scale (e.g. entire watershed or region)
Improved generality and more confidence in detected climate effects. Ignores ecological complexity and differential responses of populations.
1. Physical changes 2. Signatures in salmon pops 3. Marine survival
Fine spatial scale (e.g. individual streams/populations)

Reveals ecological complexity and differential responses of populations.
Reduced generality and little predictive power.
6. Resilience
Climate cycles on top of long-term warming

Above normal pressure over Tahiti; Below normal pressure over Australia
Below normal pressure over Tahiti; Above normal pressure over Australia
La Nia
El Nio
Source: http://en.wikipedia.org/wiki/El_Nio
Image: www.speakupforblue.com
Warm water pool; sea level well above normal
Source: http://sealevel.jpl.nasa.gov/elnino/971205.html
Its worth reflecting for a moment on what is meant by the phrase adapt to a changing environment. Intuitively, the word adapt here would seem to imply the ability of salmon to persist in a changing world, but in fact the word adaptation means something rather subtly different to an evolutionary biologist. Strictly speaking, it refers to the process whereby natural selection drives genetic changes within a population, such that genetic variants that are better fitted to the prevailing conditions increase in frequency at the expense of other, poorer adapted types. We can also broaden the definition and define adaptation at the phenotypic level, to encompass any changes in the outward characteristics of individuals such as their morphology or their behaviour, or in their internal physiological workings or patterns of development, that enhance their survival and reproductive success, or in evolutionary terms, their fitness. Such changes could be caused by underlying genetic changes or by phenotypic plasticity. But adaptation is not the same thing as persistence. A population may evolve (i.e. adapt) in response to changing environmental conditions, but go extinct in the process. At a higher level, salmon as a species or stock complexes may not go extinct, despite the loss of certain component populations. Resilience, biocomplexity etc. Genetic diversity of salmon can be considered at different levels, e.g. between stock complexes, among populations within stock complexes, and within populations. There are benefits to conserving diversity at each of these levels, but the benefits derive from quite different biological mechanisms. Within population genetic variation: grist for natural selection to do its thing. Between population/river genetic variation: increases biocomplexity, and biocomplexity increases resilience.
Peter Lisi
Spawning day of year Average Stream Temperature oC
Schindler et al. 2010
High intensity stocking might lead to loss of biocomplexity

River A River B
HATCHERY STOCK

River A River B
HATCHERY STOCK

River A River B
HATCHERY STOCK
Some studies have shown genetic homogenization effects of stocking:

Allyon et al. (2005) showed that intensive stocking of southern European rivers with Atlantic salmon of Northern European origin led to a loss of regional population structure: Prior to stocking, there were significant genetic differences (neutral loci) among neighbouring Spanish and French rivers, but these differences disappeared after stocking. Jonsson et al. 2003 found that hatchery-reared Atlantic salmon leaving the River Imsa (Norway) as smolts were more than twice as likely to stray compared with wild conspecics
Vasemgi et al. (2005) showed that extensive immigration of hatchery fish from one Baltic river into a neighbouring river with a wild population has homogenised the genetic structure (wild fish have become more like the hatchery fish)
will stocking also lead to loss of bio-complexity i.e. life history diversity and population-specific local adaptations??

Tom Reed (University College, Cork) Stocking and Climate Change

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Tom Reed (University College, Cork) Stocking and Climate Change

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Droits d'auteur :

Salmon stocking and climate change

Global scale: atmospheric CO2 is increasing

2. Signatures in salmon pops

5. Burrishoole case study

and the oceans are getting more acidic.

Global scale: average temperatures rising at unprecedented rate

Source: Moritz and Agudo Science 2013

2. Signatures in salmon pops

5. Burrishoole case study

Temperature difference from baseline global mean

Pacific decadal oscillation

2. Signatures in salmon pops

5. Burrishoole case study

Mantua et al. Bull Am Met Soc 1997

Climate change impacts on salmon

Changes in predation. Increased disease risk and parasites.

Problems due to changes in ocean climate

Source: Friedland et al. 2013 In Press

Source: Friedland et al. 2013 In Press

What about freshwater impacts on regional/local scales?

2. Signatures in salmon pops

Source: NASCO Salmon Summit report

2. Signatures in salmon pops

5. Burrishoole case study

Resilience in the face of uncertainty

ECOSYSTEM PRODUCTIVITY STOCK COMPLEX or METAPOPULATION PRODUCTIVITY POPULATION PRODUCTIVITY

Rate of generation of biomass.

Recruits per spawner Abundance

5. Burrishoole case study

Degree of local adaptation

Degree of local adaptation

Burrishoole Case study:

Longterm experiment (Burrishoole River)

to smolt survival in Burrishoole survival)

survival) survival) survival)

Winter temperature for smolts (+

Interaction between % hatchery eggs and winter temperature (-

McGinnity et al. (2009). PRSB , 276:36013610

Observed changes in climate

McGinnity et al. (2009). PRSB , 276:36013610

McGinnity et al. (2009). PRSB , 276:36013610

McGinnity et al. (2009). PRSB , 276:36013610

Resilience of stock complexes

2. Signatures in salmon pops

5. Burrishoole case study

Sockeye Salmon habitat in Bristol Bay

North Pacific Ocean

each with many populations

Salmon biological features are adapted to local

habitat conditions and how these filter climate

Salmon landscapes are shifting mosaics of suitable habitat

Number of sockeye salmon caught in Bristol Bay (1893-2010)

Ugashik Egegik Nushagak Naknek-Kvichak

catch Commercial catch Commercial

35,000,000 30,000,000 25,000,000 20,000,000 15,000,000 10,000,000 5,000,000 0

data from ADFG

Variability in salmon increases the rate of fisheries closures

Bristol Bay (intact portfolio)

Fisheries are closed 4 years every century

Number of returning salmon (millions)

average minimum spawners

Bristol Bay (eroded portfolio)

Fisheries are closed 40 times every century

Time (100 years)

Stability and productivity derive from diverse and changing habitat

Are humans causing current climate change?

Have human activities led to a net emission of CO2?