Fall 2017 A Global Issue Huge extent 40 of continental area Generally low rainfall seasonal or persistent dryness governs plantsoil relations Significant human dependence and vulnerability drought ID: 675840
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Slide1
Arid Ecohydrology
Ecohydrology
Fall
2017Slide2
A Global Issue…
Huge extent (40% of continental area)
Generally low rainfall (seasonal or persistent dryness governs plant-soil relations)
Significant human dependence and vulnerability (drought)Slide3
A Class Unto Itself…
Deep dependence on soil moisture
Soil physics
FirePlant adaptation & competition
N availabilityRoot distributions
Regular pattern developmentFeedbacks with animals (e.g., fairy rings)Slide4
Ecosystem RUE
All terrestrial ecosystems use MAP to create NPP, and are therefore, at some level, water
dependent
Differential sensitivity to variance in MAP
NPP:MAP defines rain use efficiency (RUE)
Sensitivity effectively measures the strength of water limitation vis-à-vis nutrient or evolutionary constraints
Huxman et al. (2004) - NatureSlide5
Convergent RUE
max
?
RUEmax generally occurs at low water availabilityThere is a convergence of
RUEmax across all biomes
RUEmax is close to
RUE
mean
for arid sites
RUEmax
is really low compared to RUEmean for humid sitesSlide6
Two Predictions
1 – NPP will be affected a lot if
PPT
is driven below the historic minimum
2 – Removal of other resource limitations will allow RUE to approach RUEmaxSlide7
Ergo
Water limitation imposes a common constraint on NPP across biomes
Ecosystems have the same
RUEmax despite large differences in MAP, physiology,
phylogenetic origin and climate historyAltering resource limitation underscores the relevance of biogeochemistry on NPP (i.e., compared with species)Slide8
Inhibited Deep Drainage Over Millenia
In dry areas (<500 mm) there is a nearly ubiquitous pore-water Cl
-
peak at 5-15 m below gradeMass balance of Cl yields 10,000-15,000 years of accumulation (without major downward flux event)
Consistent with ~ 1 mm/yr deep drainage
Seyfried et al. (2005) - EcologySlide9
But…
Upward water potential gradient
Water is moving out of the soil
Ergo, no downward movement
Seyfried et al. (2005) - EcologySlide10
Strong Climatic Control
Cl inventory suggests 7,000-8,000 years of accumulation at a site with 360 mm MAP
Inventory is 1,000 years at 400 mm MAP at nearby siteSlide11
Ecohydrologic Mechanism
Walvoord
et al. (2002) propose a conceptual model to reconcile these attributes
Low water potentials at the base of the root zone were established at the end of the Pleistocene (dramatically increased dryness)
Potentials (< -1 MPa) were maintained continuously during the Holocene
Any deep wetting event would reset the chloride concentrations
Slow upward movement of
water above ca. 20 m
Drainage below 20 m of Pleistocene aged water
Simultaneous recharge and upward
flux above 20 mSlide12
Vegetation is Essential
Plants are required to impose and maintain the low potentials
Two key attributes:
Deep roots (2-3 m) of sufficient density to capture all downward percolating water
Able to maintain low water potentials at depth continuouslyThe demand for continuous maintenance has enormous implications for contemporary vegetation management
Clearing vegetation can result in huge fluxes of Cl (and nitrate!) to the deep groundwaterSlide13
Deep Roots
Root density decreases exponentially with depth, though slower in arid biomes
Max rooting depths ~ 5 m, almost all co-dominant shrubs ~ 1.8 m
Root depths increased with MAP and soil coarseness
Maintaining root potentials demands cavitation avoidanceGeometry of xylem
Soil hydraulic conductance goes to 0SW USA co-dominants can withstand cavitation (embolism) from -4 to -9
Mpa
(
extremely
low)
The conceptual model requires sustained -1 MPa at the root zone
Sperry et al. (2002) – Functional EcologySlide14
Hydrologic Buffering
Arid lands are hydrologically variable
Model requires constancy
As such, moisture conditions at 2-3 m below grade must be buffered from the surface variability
Water budget suggests that Dd (deep drainage) occurs only when I > (A
ET + ΔS)
Δ
S
max
is ~ 350 mm (over 3 m root depth)Dd occurs only when I is 350 mm > A
ET (basically never)Arid land plants use
excess water in wet yearsMultiple wet years create rapid vegetation adjustmentsExpansion of deep rooted shrubsExplosion of annuals
As such, the base of the root zone basically are continuously water starved
So why invest in deep roots?Slide15
Deep Roots
C expenditure that needs to be net benefit
Existing profile data suggest very little plant available water below 60 cm
31
yr data record at 2 wk interval
Living roots at 180 cmLow potentials (< -1.5 Mpa)
Net energy benefit?
Plants may actually be hydrologically subsidizing deep roots to maintain their viability during wet periods
Confers benefits during periods of deep drought
Seyfried et al. (2001) – WRRSlide16
Implications
Natural deep rooted perennial communities are most frequently replaced with shallow rooted annuals
Grazing, planting
Changes episodic downward fluxes of water and therefore salt, nutrients, hazardous materialsExtremely difficult to reestablish deep rooted vegetationAdaptive significance of this “ecological inheritance”
Low water potentials, high Cl peaksSlide17
Alternative Stable StatesSlide18
What is a “State”
A state is a configuration of a community of organisms that can be described by a set of (dynamic) “state” variables
Biomass, density, species (ages, guilds, abundances), nutrients/organic matter
A “stable state” is a particular configuration that possesses self-reinforcing feedbacks that confer some resilience in the presence of perturbationsSlide19
Origins of the Idea
Stability of populations (constant environment)
“If the system of equations describing the transformation of state is nonlinear…there may be multiple stable points with all species present so that local stability does not imply global stability”
(
Lewontin 1969)
Overfishing, invasions, predator removalStability of ecosystems (variable environment)
(Lord) Robert May (1974)Slide20
Stable and Non-Stable Equilibria
dP
/
dt
= B(P) – D(P)
D = a*P + c
B = q + K/(1+e
-rP
)
Solve for when
dP
/dt = 0or B(P) = D(P)Slide21
The Demographic Transition
Introduces ideas:
Equilibria
Disequilibrated systemsTime lags between statesPath dependency?Slide22
Demographic Stability – Allee Effects
Per capita birth rates decline at low density
Mate finding, dispersal
Creates two population equilibriaWhich is stable? Unstable?Slide23
Balls-in-cups
Ball represents an ecosystem state
Cup represents the domain (or basin) of attraction
Remember that there are feedbacks that create “resilience”
Regime shifts occur by:Perturbation to the variables (i.e., move the ball)
Perturbation to the “fitness landscape” (i.e., change the basin)Slide24
Conceptual Model – Stability LandscapesSlide25
Ecosystem Response to Change
a. One equilibrium for each environmental condition
b. One equilibrium, but rapid rates of change over short span of conditions
c. Three equilibria can persist for one state (two stable, one unstable), strong
hysteresis (path dependency, temporal lags) in the transition between statesSlide26
Resilience – A Related Concept
Two aspects of the basin of attraction matter
Width of stability domain [Over what range of conditions/perturbations will the community recover?]
Steepness of the stability domain [How quickly will the system recover?]
Resilience can be used to define both properties The former is dubbed “ecological resilience” the latter “engineering resilience”Basically some measure of a systems capacity to buffer the effects of exogenous changesSlide27
Hysteresis – A Key Emergent Property
A systems trajectory over time may depend on its current and historical position
Changes in system configuration may delay or even prevent recovery
Suding et al. (2005)
Trends in Ecology and EvolutionSlide28
Why Should We Care?
Costly surprises (catastrophic shifts, weak ecosystem responses to management)
Collapse of fishery stocks
Outbreaks of disease
Exotic species invasionsEcosystem shifts and loss of services
Underscore the complex, contingent (dependent of antecedent conditions) nature of environmental systemsThresholds imply the need for caution when dealing with complex systems
Climate change, biodiversity loss, stock markets, land use changeSlide29
Tidal Marsh Alternative EquilibriaSlide30
Example: Shallow Lakes
Submerged Aquatics
Rooted plants little phytoplankton
Low turbidity due to:Plant filtrationHigh sediment cohesion limits resuspension due to wind action
Algal-dominatedPhytoplankton shades SAVHigh turbidity
Reduced filtrationLoose, flocculent sediments that can be easily wind-entrainedSlide31
Empirical Evidence
Enrichment of P leads to increased production
Eventually there is sufficient water column P to start to erode the controls exerted by SAV
Catastrophic flip (short time) to phytoplankton dominance
Reductions in P necessary to reverse the process are much largerSlide32
Incipient Shifts – Where are the Tipping Points?
Key problem:
It would be nice to know when these are about to happen
Interesting theoretical wrinkles:
Are the conditions for incipient regime shift the same?Volatility? Stability?
Of sufficient interest to be the subject of an NPR pieceSlide33
What Kinds of Incipient Shifts?
Multiple settings
Medicine: Onset of asthma and epileptic seizures
Finance: Stock market crashesAgriculture: Drought-society Environment: Ocean circulation, fish stocks, rangeland woody cover, boreal climate feedbacks
“Canaries in the coal mine” – predicted catastrophic bifurcationsVolatility/varianceSkewnessSlide34
Critical Slowing
At critical thresholds, small changes in state or conditions can induce a dramatic shift
That is, the system becomes increasingly “slow” in recovering
Scheffer et al. (2009)
NatureSlide35
How Do You Measure Slowing Down?
Response to experimental perturbations
Measure the rate of return
Impossible to control in large systems, but these are always perturbed
Measure rate at which system fluctuates around the meanAre the fluctuations autocorrelated
?Are they increasing in time?Are they changing in their statistical properties?
Scheffer et al. (2009)
NatureSlide36
Autocorrelation, Variance and Skewness
Guttal and Jayaprakash 2008
Ecology Letters
Scheffer et al. (2009)
NatureSlide37
Scheffer
et al. (2009)
NatureSlide38
Confronting Theory With Data
Detecting alternative attractors (observational)
Time series shifts
Bimodality in system states (across systems)
Dual relationships to a given control factor
Scheffer and Carpenter (2003)
TREESlide39
Confronting Theory with Data
Experimentally
Different initial conditions lead to different final states
Disturbance triggered transitions
Path dependency (hysteresis)
Scheffer
and Carpenter (2003)
TREESlide40
Alt. Stable State – Desert Streams
Wetlands used to be a major component of desert streams (Sycamore Creek, AZ)
Reduced to zero over the late 19
th and 20th C.
Recently become reestablished with controls on grazingSlide41
Basic Mechanism
Wetland plants control sediment dynamics
Density dependent stabilization
Floods and drying are the major hydrologic disturbance in deserts
This makes areas with dense vegetation better able to persist through floods and dry periods, and areas without vegetation less ableTwo predicted states – vegetated and gravel bed
Region of global bi-satbility defined by:
G – veg growth
S – vegetation mortality
K
s
– channel sediment stability w/o veg
C
s
– per capita stabilization of sediment
r
s
– scour vegetation mortality
V – vegetation
Q – flood frequencySlide42
Empirical Support
Water and Vegetation cover over time
Major Resetting FloodSlide43
Vegetation Effects on Vegetation Loss
(the
biogeomorphic
feedback)Vegetation density affects cover loss
Stream sites self-organize into 1 of 2 modesSlide44
Idle Restoration Thoughts
Two strategies for restoration
Change conditions (e.g., reduce P)
Force state (e.g., massive disturbance)Are there other alternative stable states lurking?Slide45
Management and A.S.S.
Native SAV
Dominated
Hydrilla
Dominated
Blue-Green
Dominated
Ecological release
Disturbance
Changes in herbivory
Nutrient enrichment
Physical removal
Chemical removal
?
Should massive disturbance be part of the restoration tool-box?
Are there states less desirable than the one we’re trying to restore?Slide46
In a Spatial Domain…(Next Time)