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Managing for Resilience in Benthic Marine EnvironmentsThe Challenge of a Sustainable Future: Contributions of LTER Science
Russell J. SchmittUC Santa BarbaraMoorea Coral Reef LTERSanta Barbara Coastal LTER
11th Annual NSF - LTER Mini-SymposiumNational Science FoundationMarch 1, 2012
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• Many ecological systems can shift suddenly from one state to another that has a reduced capacity to provide ecosystem services
• Growing concern that some abrupt shifts to alternate states may be difficult to reverse (‘regime shifts’, ‘phase shifts’, ‘ecological catastrophes’)
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• Major challenges to sustainability science & action: - forecast abrupt state changes - understand reversibility of a state change - identify means to prevent undesired shifts
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Seminal contributions of LTER science Seminal contributions of LTER science to understanding state change to understanding state change
in ecological systemsin ecological systems
• Evaluating the nature of shifts
• Developing & testing methods to warn of abrupt shifts
• Evaluating reversibility
• Identifying factors that strengthen resilience
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Time
Ecol
ogic
al s
tate
Modes of Abrupt State Shifts Modes of Abrupt State Shifts
Environmental conditions
Ecol
ogic
al s
tate
A
B
Linear Tracking
Non-linear Tracking
Hysteresis(most worrisome, most uncertain)
Coral dominates
Macroalgae dominates
A B
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Non-linear responses to gradual change in environmental conditions
Non-linear Tracking Hysteresis
Environmental conditions
B
A
Eco
logi
cal s
tate
Environmental conditionsE
colo
gica
l sta
te
A
B
adapted from Scheffer et al. 2001 Nature (North Temperate Lakes LTER)
Both cases: small change in environmental conditions can result in a large, abrupt shift in ecological state
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Two reasons to distinguish between these 2 non-linear modes
1) Reversibility of the state change
Non-linear Tracking Hysteresis
Environmental conditions
A
Eco
logi
cal s
tate
Environmental conditionsE
colo
gica
l sta
te
A
B B
• Hysteresis makes a state shift difficult to reverse – requires much greater relaxation in environmental conditions
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Non-linear Tracking Hysteresis
Environmental conditions
B
A
Eco
logi
cal s
tate
Environmental conditionsE
colo
gica
l sta
te
A
B
Two reasons to distinguish between these 2 non-linear modes
2) Respond differently to disturbance
• Non-linear Tracking – system will tend to return to pre-disturbed state
• Hysteresis – large disturbance can trigger a persistent shift to an alternate state with no change in environmental conditions
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Photo: Mike DeVries
North Temperate Lakes LTER: Persistent ‘clear-water’ (macrophyte) and ‘cloudy-water’ (phytoplankton) states in lakes
Carpenter et al. 1999 Ecol. App.
Scheffer & Carpenter 2003. TREE
Folke et al. 2004. Annual Rev. Ecol. Evol. Syst.
NTL experiment showing that a disturbance can cause a persistent shift from the ‘Cloudy-water’ to the ‘Clear-water’ state in the same environmental conditions.
NORTH TEMPERATE LAKESLong Term Ecological Research
LTER has advanced general understanding of state shifts throughtheoretical & empirical approaches
Hysteresis well described in models but challenging to find support for in nature
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Cross-site analysis of LTER time series data from 4 sites to:
1) Assess analytically whether & when abrupt transitions occurred
2) Evaluate proposed methods of forecasting abrupt shifts
3) Seek evidence of hysteresis
LTER long term data also has advanced general understanding of state shifts
California Current Ecosystem LTER
Palmer Station Antarctica LTER
Santa Barbara Coastal LTER
JRN LTERJornada Basin
LTER
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Response: Abundance of krill (Nyctiphanes simplex) - 48 year time series
Driver: Decadal climate variability in the Pacific (Pacific Decadal Oscillation - PDO)
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Response: Abundance of 3 species of penguins (Pygoscelis spp.) – 40 years
Driver: Annual duration of sea ice
AdélieGentooChinstrap
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Response: Abundance of a sea cucumber (Pachythyone rubra) – 28 years
Driver: Annual storm wave regime
P. rubra
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Jornada Basin Long Term Ecological Research
Response: Production of black grama grass (Bouteloua eriopoda) – 34 years
Driver: Growing season rainfall regime
Jornada Basin
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Issue 1: Analytical evidence for abrupt transitions in biological state Statistical breakpoints in detrended data
Santa Barbara Coastal LTER:2 abrupt transitions detected
Abrupt transition(s) identified with correct timing?
YES
YES
YES
JRN LTER
YES
Sea Cucumber
Krill
Penguins
Black Grama Grass
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Issue 2: Ability to forecast abrupt transitions
Variance increase detected?
YES ?
NO
NO
JRN LTER
NO
Models & lake studies (NTL LTER) - increased variance in time series of biological responses foreshadow abrupt transitions
(Scheffer et al. 2009 Nature; Carpenter & Brock 2006 Ecology Letters; Carpenter et al. 2011 Science)
Black Grama Grass
Sea Cucumber
Krill
Penguins
Jornada Basin LTER:Variance increased
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Issue 3: Evidence for mode of the abrupt transition (e.g., Hysteresis)Examined relationships between response & driver variables
Likely mode
JRN LTER
Sea Cucumber
Penguins
Black Grama Grass
Krill
Hysteresis mechanism? (identified by LTER studies)
Positive predation- mediated feedback
Positive feedback between soil erosion & low grass cover
Positive predator- & breeding-mediated feedbacks
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Analysis of LTER long term data
• Applying a suite of general concepts & methods to long term data advances understanding of state changes • Forecasting a shift from increased variance may have limited utility - need more frequent measures of abundance (auto-correlation)
• Analysis of driver – response relationships provides insight on the mode of response
• State changes with hysteresis appear to be occurring in multiple types of ecosystems
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Can we manage state changes?What coral reefs are telling us
• Direct control not feasible for many conditions that drive state change
• Human activities can make state change more likely; some activities highly amenable to control
• Managing for resilience: - maintain capacity of ecosystem to absorb stresses without changing
state - speed return to the desired state if disturbed
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Coral reefs frequent experience large disturbances that kill coral on landscape scales
Tropical cyclones
Outbreaks of coral predators Crown-of-Thorns Seastars (COTS)
Bleaching events
Disturbances have triggered phase shifts on some coral reefs in recent decades
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Disturbance
Disturbed reefs can return to coral dominance or switch to dominance by macroalgae
Hysteresis
Conditions
Coral A
BMacro-algae
X
Once established, macro-algae can kill adult coral & prevent coral recruitment
Herbivorous parrotfish
For reefs to be resilient, herbivores must control macroalgae
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Coral reefs of Moorea ideal for determining the features of herbivory that confer resilience
UCB Gump UCB Gump Research StationResearch Station
UCB Gump UCB Gump Research StationResearch Station
UCB Gump UCB Gump Research StationResearch Station
Why do some reefs return to the coral state while others shift to a macroalgal state?
Fundamentally related to herbivory
Moorea, French Polynesia
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• Multiple disturbances on the fore reef
• Fore reef returned to coral dominance within ~ 1 decade
• No transition to a macroalgal state
Adjeroud et al. 2009 Coral Reefs(MCR LTER – CRIOBE)
Crown-of-thorns outbreak
Bleaching event
Cyclone
Coral dynamics on the fore reef of Moorea: 1979 - 2005
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What makes the fore reef of Moorea resilient?Why haven’t disturbances triggered a shift to macroalgae?
fore reef of Moorea when MCR was established in 2004
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Providing opportunity to explore features that make the fore reef resilient
Moorea
Outbreak of crown-of-thorns seastars (COTS) Cyclone Oli
2007-2009 February 2010
Since 2004, the fore reef of Moorea experienced two more disturbances
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MCR collects time series data from 3 coral reef habitats(Fore reef, Back reef, Fringing reef) all around the island
500 mFore reef
Back reef
Fringing reef
Lagoon
Reef crest
Community dynamics differ between fore reef & lagoon habitats
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2005
2006
2007
2008
2009
2010
2011
CO
TS
den
sity
(no
. 250
m -2
)
0
2
4
6
8
2005
2006
2007
2008
2009
2010
2011
2005
2006
2007
2008
2009
2010
2011
Fore reef Back reef Fringing reef
Crown-of-thorns
Changes on the reef since 2005Lagoon
Adam et al. 2011 PLoS One
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2005
2006
2007
2008
2009
2010
2011
2005
2006
2007
2008
2009
2010
2011
Ben
thic
cov
er (
%)
0
20
40
60
80
100
2005
2006
2007
2008
2009
2010
2011
CO
TS
den
sity
(n
o. 2
50 m
-2)
0
2
4
6
8Fore reef Back reef Fringing reef
Crown-of-thorns
Coral
Changes on the reef since 2005Lagoon
Adam et al. 2011 PLoS One
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2005
2006
2007
2008
2009
2010
2011
2005
2006
2007
2008
2009
2010
2011
Ben
thic
cov
er (
%)
0
20
40
60
80
100
2005
2006
2007
2008
2009
2010
2011
CO
TS
den
sity
(n
o. 2
50 m
-2)
0
2
4
6
8Fore reef Back reef Fringing reef
Crown-of-thorns
Turfing algae
Coral
Changes on the reef since 2005Lagoon
Adam et al. 2011 PLoS One
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2005
2006
2007
2008
2009
2010
2011
2005
2006
2007
2008
2009
2010
2011
Ben
thic
cov
er (
%)
0
20
40
60
80
100
2005
2006
2007
2008
2009
2010
2011
CO
TS
den
sity
(n
o. 2
50 m
-2)
0
2
4
6
8Fore reef Back reef Fringing reef
Crown-of-thorns
Turfing algae
Coral
Macroalgae
Changes on the reef since 2005Lagoon
Adam et al. 2011 PLoS One
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2005
2006
2007
2008
2009
2010
2011
Ben
thic
cov
er (
%)
0
20
40
60
80
10020
05
2006
2007
2008
2009
2010
2011
Her
bivo
re b
iom
ass
(g m
-2)
0
10
20
30
40
50
60
2005
2006
2007
2008
2009
2010
2011
CO
TS
den
sity
(n
o. 2
50 m
-2)
0
2
4
6
8Fore reef Back reef Fringing reef
Crown-of-thorns
Turfing algae
Coral
Macroalgae
Fish
Changes on the reef since 2005Lagoon
Adam et al. 2011 PLoS One
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An MCR long term experiment has revealed:
• Macroalgae would have dominated the fore reef without control by herbivory
• Control was exerted by fishes, primarily parrotfish
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Parrotfish – 6-fold increase in biomass on the fore reef in a yearresulted from:
• Doubling in average biomass of an individual
• Increase in parrotfish abundance (numerical response)(~ 50% increase in number of parrotfish island-wide)
Parrotfish Density on Fore Reef
Tot
al le
ngth
(m
m)
100
120
140
160
180
2005
2006
2007
2008
2009
2010
Parrotfish Body Size
140
40
20
60
80
100
120
0
Bod
y B
iom
ass
(gm
s)(m
edia
n, 2
5th &
75th
per
cent
iles)
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Forereef
Lagoon
Nursery habitat in lagoon critical to numerical response of adult parrotfish on fore reef
Juvenile parrotfish nursery habitat
• Juvenile parrotfish recruit to mounding coral that only occurs in the lagoon
• Parrotfish move to fore reef within a year as they grow – a connectivity critical to resilience
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Cora
l Cov
er (%
)Co
ral C
over
(%) Fore reef Back reef Fringing reef
Lagoon
Stable nursery habitat in lagoon enabled rapid numerical response of parrotfish
Disturbances that kill coral on the fore reef do not harm parrotfish nursery habitat
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What makes the fore reef of Moorea resilient?
The capacity of parrotfish populations to respond rapidly to increases in food following the sudden loss of coral
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0.2
0.4
0.6
0.8
1.0
Parrotfish Settlement Rate
Ad
ult
par
rotf
ish
dea
th r
ate
Algae-dominatedcommunity
Coral-dominatedcommunity
Incr
ease
d fis
hing
Destruction of nursery habitat
0 10 20 30 40 50
Resilience models for Moorea show that if parrotfish grazing falls below a critical level, system undergoes phase shift from coral to macroalgae
• Hysteresis in response means grazing must be restored to a higher rate than the critical tipping point for system to return to coral
• Human activities reduce grazing in 2 ways: fishing adult parrotfish and destroying parrotfish nursery habitat.
• Grazing in Moorea still well right of the tipping point
Co
ral
Co
ver
(%) 60
40
20
0
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Managing for resilience requires ecosystem-based strategies that protect critical functions, which may be sensitive to different stresses
Artisanal fishery for parrotfish on Moorea
• Parrotfish nursery habitat sensitive to land use practices
• Fishing a key process that can trigger a phase shift
Pineapple plantation on Moorea
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The Challenge of a Sustainable Future
• Long term research critical toLong term research critical to - our understanding of abrupt state changes in ecological systems- our understanding of abrupt state changes in ecological systems
- science needed to develop sustainable management strategies- science needed to develop sustainable management strategies
Thank youThank you
• LTER has been a leader in providing new answers LTER has been a leader in providing new answers - uniquely able to combine long term data with process studies of- uniquely able to combine long term data with process studies of underlying mechanisms underlying mechanisms
- focus on long term phenomena and landscape scales necessary to- focus on long term phenomena and landscape scales necessary to reveal processes critical for resilience reveal processes critical for resilience
- network of diverse ecosystem enables comparative studies that- network of diverse ecosystem enables comparative studies that reveal generality reveal generality