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Ecosystem Succession:Ecosystem Succession:Who/What is Where and When
Dennis BaldocchiESPM
U i it f C lif i B k l
ESPM 111 Ecosystem Ecology
University of California, Berkeley
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Succession
• From the Latin, succedere, to follow after • Orderly process of community development that is
directional and predictable• Results from the modification of physical environment by• Results from the modification of physical environment by
the community– Succession is community-controlled even though the physical
environment determines the pattern rate of change and limitsenvironment determines the pattern, rate of change and limits• Culminates in a stabilized ecosystem in which biomass
and symbiotic function between organisms are maintained per unity of available energy flowmaintained per unity of available energy flow– Eugene P Odum, 1969, Science
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SuccessionSuccession
• Primary Successiony– After severe disturbance that remove or bury products of the
ecosystem• Secondary Successiony
– After disturbance on a vegetated site. Most above ground live biomass may be disturbed but soil organic matter and plant propagules remain
• Gap Phase Succession– Mortality and Tree fall for gap in canopy for new vegetation to
invade and establish itself
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Dynamic Sequence of VegetationDynamic Sequence of Vegetation
• Initial Conditions– Equilibrium
• Disturbance• Colonization/Recruitment• Colonization/Recruitment• Recovery• Competition• Succession
– Primary– SecondarySecondary– Gap Succession
• ClimaxNe Eq ilibri m
ESPM 111 Ecosystem Ecology
– New Equilibrium
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DisturbanceDisturbance• Relatively Discrete event, in time and space, that alters y p
the structure of populations, communities and ecosystems and causes changes in resource availability and the physical environment.p y
Chapin et al.
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Examples of Natural and Human-Induced Disturbance
• NaturalMortality– Mortality
• Age, Density, Self-Thinning– TreeFall– WildFire– Volcano
Fl di
Mt St Helens
– Flooding– Hurricane/Tornadoes– Insects/Disease– WindThrow– Tsunami
Logging
– Landslides– Glaciers– Sea-level Rise or Retreat
• Human-InducedL i– Logging
– Plowing– Mining– Dam Removal– Fire/Flooding
Redwood treefall
ESPM 111 Ecosystem Ecology
g– Nuclear blast/Warfare (agent Orange)
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Disturbance and Succession-Typeyp
• Primary • SecondaryPrimary– Volcano– Landslide
Secondary– Fire– Hurricane
– Flooding– Dune Formation
– Logging
– Lake Drainage– Tsunami
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Attributes of DisturbanceAttributes of Disturbance
• TypeType• Severity, Intensity, Extent
F Ti i• Frequency, Timing
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Classic Disturbance Cases in US
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Turner, 1989, Ann Rev Ecology
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Intensity and Extent
ESPM 111 Ecosystem Ecology
Foster et al 1998 Ecosystems
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ESPM 111 Ecosystem Ecology
http://www.fs.fed.us/rm/pubs/rmrs_gtr87/rmrs_gtr87_pg38.jpg
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Primary SuccessionPrimary Succession
• The process of biological colonization and change in systems that have:– no live plant material and no propagules– where the products of ecosystem processes are
either removed or buried so that there is little or noeither removed or buried so that there is little or no organic matter or organisms
• The Land is a Clean-Slate, there is no Soil.
ESPM 111 Ecosystem Ecology
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Mt St. Helens
ESPM 111 Ecosystem EcologyPhoto: ddb, 2002
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Christmas 2004 Indian Ocean TsunamiC st as 00 d a Ocea su a
ESPM 111 Ecosystem EcologyPhoto: ddb, July, 2006, Chennai, India
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Northern Example of Primary Successiono t e a p e o a y Success o
ESPM 111 Ecosystem Ecology
http://www.life.uiuc.edu/bio100
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Features of Primary SuccessionFeatures of Primary Succession
• Dispersal – getting to the siteDispersal getting to the site– Small seeds arrive first, wind blown
• Colonization and RecruitmentColonization and Recruitment– N Fixers dominate and facilitate the environment– Recruitment Success, f(soil moisture, temperature,
competition (light/soil moisture), herbivory/predation)• Establishment, Facilitation and Inhibition
– making due with the resources you can find or make
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Succession after Glacial Retreat
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Changes in Soil Properties during 1o Succession
Chapin et al., 1994, Ecol Monograph
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Secondary Succession
• Follows disturbance of an i i i hexisting community that
removes or damages the vegetation, but does not remove, destroy, or cover theremove, destroy, or cover the soil.
• Starts WITH SOIL. • PIONEER PLANTS of
secondary succession start from roots or seeds remaining in the soil or from seeds carried in by wind or animalscarried in by wind or animals from surrounding communities.
• Faster than primary succession
ESPM 111 Ecosystem Ecology
succession
http://www.life.uiuc.edu/bio100
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Start of Secondary SuccessionStart of Secondary Succession
Snags, after Yellowstone Forest Fire
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Example of Secondary Succession
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Features of 2nd SuccessionFeatures of 2 Succession
• Colonization– Sprouting from root and stems– Germination from soil seed bank– Advanced regeneration by long-lived shade tolerant– Advanced regeneration by long-lived, shade tolerant
seedlings– Dispersal and In-fill from adjacent gaps
• Wind blown seeds Birds Animals• Wind blown seeds, Birds, Animals• Facilitation
– Hydraulic Lift– Shade and protection of seedlings from desiccation
• Competition• Herbivory and Pathogens
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• Herbivory and Pathogens
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Long Distance Dispersal of Plants
ESPM 111 Ecosystem EcologyNathan 2006, Science
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Conceptual Paradigm: NEP is a function of age
ESPM 111 Ecosystem EcologyOdum, 1969, Science
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Roles of Climate, Forest Type and Succession Type on Disturbance on NEP
NEP
TIME
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Stand Carbon Dynamics and Disturbance
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Ecosystem Dynamics, CasesFireLoggingTreefallTreefallOld Field Succession
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The boreal forest is dominated by fire
Large fires1980-99
ESPM 111 Ecosystem EcologyCourtesy of Brian Amiro, Univ Manitoba
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Combustion lossesCO2, CO, CH4
Decomposition
Decomposition Renewed CWD,
regenerationRenewed mature forest stand
Successional vegetation to crown
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vegetation to crown closure
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Successional trajectory (Goulden et al. Northern M nit b Bl ck Spruc chr n s qu nc )Manitoba Black Spruce chronosequence)
1989 burn
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Role of Repeated Surface Fires
ESPM 111 Ecosystem EcologySchulze et al 2000, Science
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Effects of Stand Age:After LoggingAfter Logging
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Law et al. 2003 Global Change Biology
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Role of N deposition on NEP and GPP Dynamics across Europe
ESPM 111 Ecosystem EcologyMagnani et al 2007 Nature
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Energy Exchange and Stand Age
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Boreal Forest
o0.20
AKMB
e A
lbed
o
0.15
MBSKSOA
mm
ertim
e
0 05
0.10
Sum
0.00
0.05
Time since disturbance (years)0 25 50 75 100 125 150 175
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Amiro et al., 2006 AgForMet
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Ave ET along a chronosequence, 2001-2005g q ,
• Abandoned old agricultural Field. 562 mm
• Successional Pinus taeda, 658 mmOld h d d f t 617• Old hardwood forest, 617 mm
Stoy et al. 2006 GCB
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y
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Regeneration after Fire: Salvage Log or Not???
Post-Fire Logging can be counter-productive to the goals of forest regeneration and fuel reduction by removing naturally seeded conifers and increase surface fuel load
ESPM 111 Ecosystem EcologyDonato et al 2005 Science
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Regeneration after Fire
Donato et al Science, 2005
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Oak Recruitment/Colonization in California
ESPM 111 Ecosystem EcologyTyler et al 2006 Quart Rev Biol
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Old-Field Succession42 year Chronosequence of abandoned vineyards in France
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Shipley et al 2006 Science
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Succession and Resources
Soil N plays a role in composition, species richness and successional dynamics
ESPM 111 Ecosystem EcologyTilman 1987 Ecol Monograph
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Community Energetics
Developmental Stage Mature Stage
G >1 1Gross Production/Respiration
>1 ~1
Gross Production/Standing
high lowProduction/Standing BiomassYield High Low
Food Chains Linear, grazing Web-like
ESPM 111 Ecosystem EcologyOdum, 1969
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Community Structure
Developmental Stage Mature Stage
T t l O i M tt S ll LTotal Organic Matter Small Large
Inorganic Nutrients extrabiotic Intra biotic
Species DiversityLow High
Growth Form Rapid growth, r selection Feedback control, K l tiselection
Niche Specialization Broad Narrow
Life Cycle Short, Simple Long, ComplexLife Cycle Short, Simple Long, Complex
ESPM 111 Ecosystem EcologyOdum, 1969
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Ecological Succession - Overviewco og ca Success o O e e
• From the Latin, succedere, to follow after• "Change in the species composition of a community over time." (Lewis, Life glossary)• Primary Succession follows the formation of new land surfaces consisting of rock, lava, volcanic
ash, sand, clay, or some other exclusively mineral substrate. – This means that there is NO SOIL present. – Soil is a mixture of mineral material, decaying organic material, and living organisms.
• Secondary Succession follows the destruction or partial destruction of the vegetation area by t f di t b lik fi i d t fl d th t l th il i t tsome sort of disturbance, like a fire, windstorm, or flood that leaves the soil intact.
• Pioneer species initiate recovery following disturbance in both primary AND secondary successions
• Pioneers "pave the way" for later colonists by altering the biotic and abiotic environment:– soil stabilization
il t i t i h t ( i tt d bi l i l it fi ti )– soil nutrient enrichment (organic matter and biological nitrogen fixation) – increased moisture holding capacity – light availability – temperature – exposure to wind
• Species composition tends towards a Climax Community through succession• Species composition tends towards a Climax Community through succession.• The climax community describes an end product of succession that persists until disturbed by
environmental change.• Succession occurs at large scales involving higher plants and animals, but may involve microbial
communities on a smaller scale.
ESPM 111 Ecosystem Ecologyhttp://www.life.uiuc.edu/bio100
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Points to PonderPoints to Ponder
• Do Ecosystems Reach Climax states?Do Ecosystems Reach Climax states?– Does the frequency of Disturbance prevent climax
states from Occurring?– When? Where??
• How does the disturbance induced time series of G f ?NEP, GPP and Reco vary with logging vs fire?
• What are the Pros and Cons of Facilitation?– e.g. recruitment of ponderosa pine seedlings under
ceanothus or bitterbrush and effects on water, light and nutrients
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ESPM 111 Ecosystem Ecology
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ESPM 111 Ecosystem EcologyOdum, 1969
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ESPM 111 Ecosystem Ecology