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Responses of NW European nearshore and
coastal ecosystems to climate change and
other impacts: combining sustained
observations, experiments and modelling
Prof Steve Hawkins
Ocean & Earth Science, National Oceanography
Centre Southampton
(Dean, Faculty of Natural and Environmental
Sciences) University of Southampton
Korea Feb 2014
Our work at the Marine Biological Association of
the UK, Bangor and now Southampton
Steve Hawkins, David Sims, Martin Genner, Mike Burrows, Nova Mieszkowska,
Alan Southward
Mike Kendall, Rebecca Leaper, F. Pannaccuilli, Georgina Budd, Patricia
Masterson, Richard Thompson, Matt Frost, Paula Moschella, Pippa Moore,
Jackie Hill, Stuart Jenkins, Louise Firth, Keith Hiscock, Elvira Poloczanska and
many others
Past and future climate change
Intergovernmental Panel on Climate Change (IPCC) 2007
Global Environmental Change
Climate:
•
•
•
Not just temperature
But also storms, precipitation,
frequency of extreme events, NAO
index (more positive years)
(Reduction of ocean pH – strictly
not climate change)
Will influence biodiversity:
•Shifts in environmental gradients (desiccation, wave action, stratification,
salinity etc.)
•Changes in frequency of disturbance events
•Poleward migration of species
•Changes in assemblage composition & interactions (often via recruitment
regimes)
•Greater likelihood of non-native invasions
Climate Change in the N.E. Atlantic
Particularly strong warming
has occurred in the North
Atlantic since the mid1980s (35° to 65°N, 0° to
35°W)
This is 2 x rate of any
previous warming event
on record, 0.5-1°C in last
20 years
 Exceeds global average
IPCC Climate Change 2001 The Scientific Basis
Local artisanal to global industrial ocean use
Cornwall 1890s
6
Outline of talk
• Introduction to MBA time-series
• Restart of off-shore time series collected at
Plymouth (lab project)
• Rocky shore indicators: Broad-scale re-surveys
and time series restart (led by SJH)
• Modelling enabling forecast and prediction? (led
by Mike Burrows)
• Policy implications – adapting to climate change
Classic experiment repeated by Hawkins 1981
Limpet removal strip
2 months
9 months
NS Jones,1946 , 1948
5 years
Hawkins, 1981 Kieler Meeres. (EMBS 1980)
Patella depressa: a ‘southern’ species higher
abundance (as% total limpets) in 1950s cf 1980s
1950s 1980-84
Not found
Rare
Occasional
Frequent (1-10%)
Common (11-50%)
Abundant (>50%)
Half symbols
indicate no
change in scale
Hawkins et al., unpublished,
Kendall et al. 2004 IBIS
P. depressa
southern
P. vulgata
northern
Plymouth sail-fishing fleet
Fisheries investigations
began in 1887 ahead of
building the lab
Stanhope A. Forbes, 1885
Mean annual sea surface temperature
1871-2006 off Plymouth
13.5
(grid square 50–51° N, 04–05° W).
Mean annual SST (ºC)
13.0
12.5
12.0
11.5
MBA lab
founded
Observed
5 yr mean
11.0
1870
1890
1910
1930
1950
1970
1990
2010
Year
Data from the UK Meteorological Office Hadley Centre. Much of it collected by
the MBA/PML
Why are the British Isles & Ireland sensitive?
Britain straddles a biogeographic boundary, resulting in species with
northern and southern biogeographic distributions co-existing.
Also multiple range limits
Forbes, 1858. A.K. Johnson’s Physical Atlas.
The western English Channel
Major long-term sampling
stations off Plymouth
Regular intertidal stations
From Southward et al., Adv. Mar. Biol., 2005
MBA Time Series: English Channel
Temperature and Salinity
E1
1902-1987
Nutrients
E1
1921-1987
Phytoplankton
E1
1903-1987
Primary production
E1
1964-1984
Zooplankton
E1, L5
1903-1987, 1995-1998
Planktonic larval fish
E1, L5
1924-1987, 1995-1998
Demersal fish
L4
1913-1986
Intertidal organisms
various 1950-1998, 1997- date
Infaunal benthos (intermittent)
L4
1922-1950
Epifaunal benthos (intermittent)
L4
1899-1986
PML time series:plankton & hydrography at L4 since 1987
n.b. There are many gaps in these series
MBA Time Series: English Channel
Western Channel Observatory from 2007 PML/
MBA/SAHFOS as part of Oceans 2025
Temperature and Salinity
E1
1902-1987, 2002Nutrients
E1
1921-1987, 2002Phytoplankton
E1
1903-1987, 2002Primary production
E1
1964-1984
Zooplankton
E1, L5
1903-1987, 1995-1998,2002Planktonic larval fish
E1, L5
1924-1987, 1995-1998,2002Demersal fish
L4
1913-1986, 2001-2003,2005Intertidal organisms
various 1950-1998, 1997/2001-2005,Infaunal benthos (intermittent)
L4
1922-1950, 2003
Epifaunal benthos (intermittent)
L4
1899-1986, 2005PML time series:plankton & hydrography at L4 since 1987
n.b. There are many gaps in these series,
Defra & Agg. levy funded restarts in red
Reviewed in Southward et al., Adv. Mar. Biol. 47:1-105, 2005
Catch (tonnes)
Landings of pelagic fish at Plymouth
8000
7000
6000
5000
Herring - Clupea harengus
4000
3000
2000
1000
0
1920 1930 1940 1950 1960 1970 1980 1990 2000
Year
Sutton Harbour, 1925
Such fluctuations occur
back to the middle-ages
(Southward et al. 1988 JMBA 68(3):
423-445)
Catch (tonnes)
6000
Pilchard - Sardina pilchardus
5000
4000
3000
2000
1000
0
1920 1930 1940 1950 1960 1970 1980 1990 2000
Year
Data source: UK Government records
pilchard eggs (monthly mean)
Pilchard eggs and non-clupeid larvae
40000
pilchard (Sardina pilchardus) eggs
10000
larval fish
8000
30000
6000
20000
4000
10000
0
1920
2000
1940
1960
Year
1980
0
2000
8000
7000
6000
5000
Herring - Clupea harengus
4000
3000
2000
1000
0
1920 1930 1940 1950 1960 1970 1980 1990 2000
Year
Sutton Harbour, 1925
6000
Catch (tonnes)
Catch (tonnes)
Landings of pelagic fish at Plymouth
Pilchard - Sardina pilchardus
5000
4000
3000
2000
1000
0
1920 1930 1940 1950 1960 1970 1980 1990 2000
Year
Data source: UK Government records
Interaction of fishing and climate
Plymouth inshore demersal fisheries
Mean catch per hour
(individuals)
Climate change
Breams (Family Sparidae)
5
4
3
2
1
0
Mean catch per hour
(individuals)
1910
1930
1950
1970
1990
2010
Rays (Family Rajidae)
16
12
8
4
0
1910
1930
1950
1970
1990
2010
Shark Trust
Fishing pressure
Genner et al., unpublished
MBA Trawls: fewer large fish
species in catches due to fishing
October 1963
November 2001
August 2006
Sharpest declines seen in large species:
skate & ray, brill, conger eel
-3
Light
1956
1955
1957
1952
1953
Increase
Principal Component
Analysis (PCA)
-6
965 1985 2005
-8
-4
0
4
8
to identify
species
responding
Warm
PC1 (Sea Temperature)
Cold
0.4
Species +vely
correlated
with more
fishing
d
Species
increasing
965
1985
2005
with
warming
Decline
PC 2 (More Fishing)
0.2
0
-0.2
-0.4
-0.4
Increase
-0.2
0
PC1 (Warming)
0.2
0.4
Species
declining
with
fishing
Decline
Genner, Sims, Wearmouth, Southall, Southward, Henderson & Hawkins, 2004, Proc. R. Soc. of Lond B
PC1, an index of community-level
change, correlates with sea temperature
r2 = 0.73; P < 0.001
Fish Community Structure
(PC Axis 1)
10
6
2
-2
-6
-10
11.5
12
12.5
13
Mean Annual SST (ºC)
Genner et al. (2004) Proc. R. Soc. London B 271: 655-661
13.5
18000
16000
13.0
12.5
12.0
11.5
Demersal Landings
500
14000
450500
400450
12000
400
350
10000
1930
1950
1970
1990
6000
4000
0
1910
2010
War years
1930
1950
Year
1970
1990
2010
Year
6
0 0
1910
1910
1930
1930
1950
1950
1970
1970
19901990 2010 2010
Year
Year
6
Large species
Small species
4
PC1 (mean monthly)
4
PC1 (mean annual)
350
300
300
250
250
200
200
150
150
100
100
50
50
8000
2000
11.0
1910
Abundance (Mean individuals per hous)
hour)
individuals
annual,
(Mean
Abundance
hous)
perper
individuals
(Mean
Abundance
Sea surface temperature
temperature
2
0
-2
2
0
-2
-4
-4
-6
-6
1910
1930
1950
1970
Year
1990
2010
1910
1930
1950
1970
Year
1990
2010
hour)
individuals
annual,
(Mean
Abundance
h ous)
ls per per
n individua
(Mea
Abun dance
13.5
Demersal fish landings (tonnes)
Mean annual sea surface temperature (
°C)
English Channel fish assemblages
Small species – follow climate change
Large species - abundance declines over the last 50 years
14
12
10
8
6
4
2
0
1910
1930
1950
1970
1990
2010
Year
Co-funded by DEFRA (projects MF0727 and MF0730; 2000-2003)
Genner et al GCB (2010)
English Channel fish sizes 1913-2006
Rays
Plaice
Small and medium species – no change
Large species – fewer large individuals
3500
12000
3000
10000
500
0
1910
1930
1950
1970
1990
2010
Year
3500
3000
2500
2000
1500
10
2000
0
1910
-10
1950
1970
1990
2010
1970
1990
2010
-20
40000
-30
35000
30000
-40
25000
-50
100
1,000
10,000
100,000
20000
15000
10000
500
0
1910
1930
Year
10
1000
6000
4000
0
Mass (g)
1500
1000
Mass (g)
Mass (g)
8000
2000
Change in mass (slope)
Length (mm)
2500
Maximum body mass of species (g)
1930
1950
1970
Year
1990
2010
5000
0
1910
1930
1950
Year
John Dory
Monkfish
Genner et al GCB (2010)
Fish populations in the English Channel
• Massive changes in assemblage composition
• Interaction of climate & fishing driving
changes in demersal fish
• Climate influenced advance of southern
species; northern persisters?
• Good evidence of climatic impacts on pelagic
species : Herring-pilchard(sardine) switches
back to 13th century (Southward et al. 1998
JMBA)
SHIFTS IN RANGE and
CHANGES IN ABUNDANCE
ON ROCKY SHORES
Hawkins et al 2009 MEPS
Intertidal organisms - inexpensive
indicators of change in nearshore ecosystems
• Easily sampled non-destructively on broad-scale
• Amenable to mapping and hierarchical statistical
approaches
• Processes can be understood from experimental
ecology
• Larval success in offshore environment reflected in
recruitment
• Good time-series availability (N & S pairs of spp)
• Not exploited (at least in UK)
MarClim resurvey locations
•Building on the Crisp and
Southward baseline from 1950s
•over 400 sites surveyed
• sites beyond range edges
• Irish team established 2003
• 70 sites surveyed
• intercalibration in N. Ireland
Marine Biodiversity and Climate Change Project:
MarClim (2001 –2005 and since)
Long-term datasets exist for temperature sensitive rocky
shore intertidal species.
Aims:
•Collate & archive data, broad-scale resurvey of 1950s sites
•Continue and re-start time-series of key species
•Demonstrate changes that have occurred in intertidal
species
•Develop and test hypotheses on climate change
•Forecast future changes based on climate scenarios using
modelling
Mean annual SST (ºC)
Sea-surface temperature
offshore Plymouth 1905-2005
13.5
13.0
12.5
12.0
11.5
11.0
1905 1925 1945 1965 1985 2005
Year
Data source: Met Office Hadley Centre
Grid square 50-51ºN, 4-5ºW
See also Sheppard, 2004, Mar. Poll. Bull., 49: 12-16
Example of changes in the distribution of a intertidal
species in Northern Scotland (spring 2002)
• northern range extension of
~70km in North Scotland since
1986 (Miezskowska, unpub)
Top shell – Gibbula umbilicalis
Changes in population dynamics near northern
range edge
% of sample
population
1983
Frequent recruitment failure in
cooler 1980s
50
40
30
20
10
0
1
3
5
7
9
Successful recruitment every
year in warmer 2000s
11 13 15 17 19
basal diameter (mm)
2004
50
40
30
20
10
0
% of sample
population
% of sample
population
2003
1
3
5
7
9
11 13 15 17 19
basal diameter (mm)
50
40
30
20
10
0
1
3
5
7
9
11 13 15 17 19
basal diameter (mm)
Mieszkowska et al. in submission, G. umbilicalis at Culkein, N. Scotland
Range edge distribution of intertidal species
in the English Channel (summer 2004-2005)
a)
d) Patella depressa
Southampton
Key:
Not found
Rare
Occasional
Frequent
Common
Abundant
N
40 km
e) Patella ulyssiponensis (P. aspera)
Southampton
b) Gibbula umbilicalis
f)
Southampton
Osilinus lineatus
Southampton
g)
c)
Southampton
Species distribution (1950s, 1980s)
Herbert,
Hawkins,
Southward,
Sheader,
Hawkins,
Herbert
&
&
Sheader
2003
Southward,
2003
Southampton
Range extension (2000-2004)
Range limit
Hawkins, Mieszkowska, Moschella new data, 2004-2005
LCS
Other coastal
defence
structures
Broad-scale
modification of
coastline
Elmer defence
scheme
Range edge distribution of intertidal species
in the English Channel (summer 2004-2006)
a)
d) Patella depressa
Southampton
Key:
Not found
Rare
Occasional
Frequent
Common
Abundant
N
40 km
e) Patella ulyssiponensis (P. aspera)
Southampton
b) Gibbula umbilicalis
f)
Southampton
Osilinus lineatus
Southampton
g)
c)
Southampton
Species distribution (1950s, 1980s)
Sheader, Hawkins,
Hawkins, Herbert
Sheader,
Herbert&&
Southward,
2003
Southward,
2003
Southampton
Range extension (2000-2004)
Range limit
Hawkins, Mieszkowska, Herbert, Moschella new data, 2006
Patella depressa: a ‘southern’ species
abundance (as% total limpets) 1950s and 1980s
1950s 1980-84
Not found
Rare
Occasional
Frequent (1-10%)
Common (11-50%)
Abundant (>50%)
Half symbols
indicate no
change in scale
Hawkins et al., unpublished,
Kendall et al. 2004 IBIS 146: 40-47
P. depressa
southern
P. vulgata.
northern
Proportion of P. depressa to total limpets
Increases in the proportion of P.
depressa a southern species out of the
total mid shore limpet population
(P.vulgata plus P.depressa)
Woolacombe
Duckpool
Hartland Quay
y = 0.008x - 15.835y = 0.0111x - 21.526 y = 0.0108x - 21.06
R2 = 0.4126
R2 = 0.5578
R2 = 0.5704
1
0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
1980
1983
1986
1989
1992
For all regression analyses P<0.001
1995
1998
2001
2004
Hawkins life time’s unpub work
Patella depressa: a ‘southern’ species abundance
(as% total limpets) 1950s and 1980s
P. depressa breeding
populations reestablished
P. depressa now
80%
60-
P. depressa now >50%
on many south coast
locations
Hawkins et al., unpublished, Kendall et al. 2004 IBIS 146: 40-47
Range extensions of southern species
Gibbula umbilicalis
Purple topshell
Chthamalus montagui
Stellate barnacle
September 2006
Osilinus lineatus
Toothed topshell
Melaraphe neritoides
Small periwinkle
Bifurcaria bifurcata
Brown alga
Balanus perforatus
Acorn barnacle
Decreases in abundance and retreats of
northern species
Tectura testudinalis
Tortoiseshell limpet
Alaria esculenta
Brown alga
Semibalanus
balanoides
Acorn barnacle
Modelling using barnacles
• Predictive modelling of interacting
species
• Statistical analysis of past data sets
• Process-based modelling including
indirect effects
• Forecasts using scenarios
Semibalanus balanoides (northern species)
Chthamalus montagui
Chthamalus stellatus
(southern species)
MBA time series: abundance of barnacles in
S.W. England (8 sites south coast)
Chthamalus spp.
HWN
Semibalanus balanoides
HWN
spp.
HWN
MTL
HWN
MTL
10
no per cm
2
8
LWN
MTL
LWN
MTL
LWN
LWN
SJH/ MarClim
AJS
Chthamalus spp.
‘Southern’
6
4
2
Semibalanus spp.
‘Northern’
0
1950 1956 1962 1968 1974 1980 1986 1992 1998 2004
1997-2003 NERC small grant
Southward (1967, 1980, 1991) then Hawkins;
Barnacle Time Series: Cellar Beach,
S.W. England
Southward, 1991, JMBA
Competition: Evidence
• Semibalanus balanoides
– Mid and low shore, high mortality on
high shore
– Single brood
– Juveniles settle in spring, grow fast
– Dominant competitor
• Chthamalids
– High shore, resistant to desiccation
stress
– Lower limit set by competition by S.
Competitive exclusion
balanoides
experimentally verified in the field
– Multiple broods
by J. Connell (1961)
– Juveniles settle in summer &
autumn
Correlation of adult abundance with monthly lagged
sea surface temperature (SST)
0.4
0.3
•
Semibalanus balanoides
– Negative correlation with
SST previous June
• High recruit mortality
in warmer years?
• Lower larval mortality
in cooler years?
•
Chthamalids
– Positive correlation with
SST previous June
• Lower juvenile
mortality in warmer
years as poor survival
of S. balanoides
juveniles?
0.2
0.1
0
-0.1
-0.3
High Shore
-0.4
Mid Shore (SB)
-0.5
Mid Shore (CM)
Low Shore
-0.6
-3
-2.5
-2
-1.5
-1
-0.5
0
0.5
1
0.6
0.4
0.2
0
-0.2
-0.4
-0.6
-3
-2.5
-2
-1.5
-1
Lag Year
-0.5
0
0.5
1
Pearson’s correlation
-0.2
Is climate acting on each species directly or is
climatic influence mediated by the presence
of a competitor?
High Shore
-0.49 *
SB
-071 ***
SST
y-1
0.17 ns
Ch
Mid Shore
-0.52 **
SB
SST
y-1
-0.72 ***
0.10 ns
Ch
Causal Path Analysis
– SST the previous June directly
influencing adult S. balanoides
abundance only
– S. balanoides abundance
influences adult Chthamalid
abundance
– The influence of climate on
Chthamalids is mediated by the
presence of S. balanoides, the
dominant competitor, in SW
England
Model Output
Hypothesis 1
Hypothesis 3
S. balanoides recruitment driven by SST
S. balanoides recruitment driven by SST
S.1.4 balanoides
2.0
1.8
1.2
Adults per cm2
Historical
abundance
S. balanoides
Interference competition between
juvenile S. balanoides and
Chthamalids
S. balanoides
1.6
1.0
1.4
1.2
0.8
1.0
0.6
0.8
0.4
0.6
0.2
0.4
0.2
0.0
1920
1940
1960
1980
0.0
1920
1940
1960
1980
Historical
abundance
chthamalids
Adults per cm2
Model output
3.5
3.5
3.0
3.0
2.5
2.5
2.0
2.0
1.5
1.5
1.0
1.0
0.5
0.5
Chthamalids
0.0
1920
Chthamalids
0.0
1940
1960
1980
1920
1940
1960
1980
Model output Hypothesis 3: 1955 - 2100
Low emissions
S. balanoides
1.6
Historical
abundance S.
balanoides
High emissions
S. balanoides
1.6
1.4
1.4
1.2
1.2
1
1
0.8
0.8
0.6
0.6
0.4
0.4
0.2
0.2
0
0
1955
1980
2005
2030
2055
2080
1955
1980
2005
2030
2055
2080
Model output
3.5
Historical
abundance
chthamalids
Chthamalids
3.5
3
3
2.5
2.5
2
2
1.5
1.5
1
1
0.5
0.5
0
1955
Chthamalids
0
1980
2005
2030
2055
2080
1955
1980
2005
2030
2055
2080
Polanczska et al: Ecology 2008 NERC grant and
Marclim project
Consequences of climate driven
change in assemblage
composition(species identity) for
ecosystem functioning – does it
matter?
Patella depressa: a ‘southern’ species
abundance (as% total limpets) 1950s and 1980s
1950s 1980-84
Not found
Rare
Occasional
Frequent (1-10%)
Common (11-50%)
Abundant (>50%)
Half symbols
indicate no
change in scale
Hawkins et al., unpublished,
Kendall et al. 2004 IBIS 146: 40-47
P. depressa
southern
P. vulgata.
northern
Limpet (P.vulgata) –Fucus –barnacle
interactions
More
Burrows & Hawkins, MEPS 1998
Differences in spatial distribution
Ring statistics O(r)
0.3
0.25
P. vulgata preferential
aggregates beneath
Fucus at scales of up to
50cm = size of Fucus
patch.
0.2
0.15
0.1
0.05
0
1
3
5
7
9
11
13
15
17
19
21 23
25
27
29
31
Spatial scale r [cells]
Ring statistics O(r)
0.03
0.025
P. depressa does not
aggregate or avoids
Fucus
0.02
0.015
0.01
0.005
0
1
3
5
7
9
11 13
15 17
19 21 23
25 27 29
31
Moore et al unpub
Spatial scale r [cells]
95% confidence intervals generated following 1,999 Monte Carlo simulations
See Wiegand and Moloney (2004) for methodology
Mean limpet mortality per
replicate (n=5)
Different behavioural responses
5
4
• Over 33% of P. vulgata individuals
died following Fucus removal.
3
2
• Fucus removal had little effect on
P. depressa mortality.
1
0
P. vulgata
Fucus removal
Fucus patch
Open rock
400
Distance (mm)
P. depressa
300
200
100
• Many P. vulgata individuals
relocated home scars often beneath
nearby Fucus patches.
•There was little response by P.
depressa, although they were less
loyal to home scars beneath Fucus
patches.
0
P. vulgata
P. depressa
Moore et al 2007 MEPS 334: 11-19
Limpet (P.vulgata) –Fucus –barnacle
interactions
More
Burrows & Hawkins, MEPS 1998
Limpet –Fucus –barnacle interactions: adding
extra species and changing abundance
More Chthamalus less
chance of algal
escapes
Less Semibalanus
More Chthamalus ,
barnacles less
dense
More
More P. depressa less
aggregation, less chance
of algal escapes
More grazer
species less
chance of
escapes
Limpet –Fucus –barnacle interactions: adding
extra species and changing abundance
More Chthamalus less
chance of algal
escapes
Less Semibalanus
More Chthamalus ,
barnacles less
dense
More
More P. depressa less
aggregation, less chance
of algal escapes
More grazer
species less
chance of
escapes
Semibalanus balanoides
Mixed
S.balanoides grows
faster hence lower
secondary
productivity with
chthamalids ??
Chthamalus montagui
Fucus escapes more
likely on dense
S.balanoides ???
Consequences of change
• Lower frequency of
Fucus escapes
• Possible shift in
balance along wave
action gradient
• Less dynamic system
due to difference in
behaviour of key limpet
species
Climate change will result in assemblages dominated by
suspension feeders and grazers
Fucoid dominated
Patchy
Barnacles / limpets
with less primary producers and export of detritus
Global change – functional
consequences?
• Balance between grazers/ suspension feeders &
fucoids will alter
• Lower probability of algal escapes from grazing
• Increased interaction of stress & grazing on
early stages
• Less shelter and lower diversity
• Fewer primary producing fucoids and less
detritus
• Likely to be rapid non-linear step change
Globalization
Harbour wall - Nelson, N.Z.
Elminius and Mytilus
S. Indopacific
Breakwater - Liverpool, U.K.
Mytilus and Elminius
N. Pacific & Atlantic
Homogenisation
Global climate change
Sea Level Rise, extreme waves & storm surges
“Hardening” of coasts
Coastal defence schemes
Environmental heterogeneity
Roughness, crevices & pools
Habitat enhancement
Drill-cored rock pools at Tywyn
© M. Gee Chapman
© M. Gee Chapman
Firth et al. (in sub)
Coastal Engineering
Habitat enhancement
The BIOBLOCK
Habitats
•Rock pools
•Horizontal crevices
•Pits
Louise Firth
now at NUI Galway
Firth et al. (in sub)
Coastal Engineering
Adaptation to climate change needs the
ability to separate low amplitude long
wave length climate driven change from
regional and local scale impacts
We can do nothing about climate change for the next 50
years ( until mitigation or new technologies kick in)
•Manage interactions of climate change with those
things we can control
•Climate with global: non-native species
•Climate with regional: fishing and eutrophication
•Climate with local: inappropriate coastal development
Hawkins, 2012 Aquatic Conservation, Hawkins et al 2013 Marine Policy,
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