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Inputs
Outputs
EcosystemProductionConsumptionDecompositionElement cycling
FEE so far
Energy flowChemical transformation
The extended physics of biota
Fundamentals of Ecosystem Ecology. Monday January 14, 2013, AM
Physical Ecosystem Engineering by Organisms
Clive G. Jones
How organisms physically alter the abiotic environment & the consequences for them, other species, other ecological processes &
interactions, & ecosystem & landscape functioning
http://www.outdoorescapesnewhampshire.com/Photos/BeaverDam.jpghttp://ci.marysville.wa.us/publicworks/swm/docs/beaver_files/Beaver%20Pic.jpg
Beaver → Dams → Hydrology, sediments → Biogeochemistry, habitat
Two of countless examples
http://library.thinkquest.org/05aug/00183/How%20Hurricanes%20Impact%20the%20Reef.htm
Coral reefs → Wave attenuation → Refugia
Xxxxx →Xxxxxx → Xxxxxx → Xxxxxxx
Outline
Ecosystem Context
Cause-Effect, Time & Space
Identity & Ecosystem Functioning
Frontiers Beyond & Within Ecology
A place with all the living & non-living interacting – Tansley 1935 Abiotic > Abiotic: Physical & chemical processese.g., erosion & deposition (material vectoring), … ; redox, precipitation, hydrolysis, …
Biotic > Biotic: Direct energy, material, information transfere.g., predator-prey, trophic mutualism, biotic resource competition, pollination, …
Abiotic > Biotic: Conditions & resources – ‘Abiotic Determinism’e.g., climate, topography, parent materials, pH, salinity, redox…
Biotic > Abiotic: Assimilation & dissimilation (uptake & ‘waste’ transfers) – Trophice.g., plant nutrient uptake & OM production, urine, feces, microbial mineralization
Physical ecosystem engineering process – Non-trophice.g., beaver dam & pond, root macropore & drainage, coral reef wave attenuation …
All Interactinge.g., nutrient cycling; direct abiotic resource competition; physical ecosystem engineering consequence (biogeochemical heterogeneity, habitat, engineer feedbacks,…); …
Controls on ecosystem structure & functioning
Also see: Weathers, K. C., Ewing, H. A., Jones, C. G., and Strayer, D. L. 2012. Controls on ecosystem structure and function. Pp 215-232 (Chpt. 11) In: Weathers K. C., Strayer D. L., and Likens, G. E. (eds). Fundamentals of Ecosystem Science. Academic Press.
ABIOTA
Physical &
Chemical Processes
Assimilation & DissimilationPhysical Ecosystem Engineering
Abiotic Determinism
BIOTA
Direct Energy,
Material, & Information
Transfers
Engineered ecosystem (patch) & landscape
Engineered: Physical control by organisms on internal structure & function
Unmodified: Not engineered by the focal engineer(s)
Landscape = n engineered + n unmodified in space
EMO: Fluxes of energy, materials, organisms
Engineered Unmodified
EMO
Outline
Ecosystem Context
Cause-Effect, Time & SpaceDefinitionsFramework: Mostly TemporalSpaceSpatio-Temporal Dynamics
Identity & Ecosystem Functioning
Frontiers Beyond & Within Ecology
Jones & Gutiérrez 2007
Definitions
PhysicalNon-Trophic
Abiotic resourcesAbiotic conditions
Biotic effect
Process
Consequence
Ecosystem
Jones et al. 2010
Framework: Mostly Temporal
Process, consequences, system dynamics
Magnitudes
Engineering Process
Engineer > structural change > abiotic change
Exemplify!
Engineer structure formation (F)
1. Necessary? Sufficient?
2. Autogenic/allogenic?
3. Constraints?
4. Density & per capita engineering activity: necessary/sufficient?
5. Structural legacies?
6. Measure? Units?
Structural decay (D)
1. Agents?
2. Why occur?
3. Determinants?
4. Offset?
5. Legacy fate?
6. Measure? Units?
Net structural change (F-D)
1. Why relevant?
2. Measure? Units?
Abiotic change
1. Why occur?
2. Determinants?
3. Measure? Units?
4. Relationship between abiotic change & structural decay?
Engineering Consequence
Structural/abiotic change > Biotic change & engineer feedbacks
Exemplify!
1. Why occur?
2. How predict? Expectations?
3. Measure? Units?
4. Why feedbacks relevant?
5. Kinds of feedback? Time scales? Consequences?
Space
EngineerEngineering requirementsStructural- & abiotic-state dependence
Other speciesEngineered habitat specialistsUnmodified habitat specialistsHabitat generalists
Energetic & material connectance
Engineered Unmodified
EMO
1. General model of environmental dynamics
Raynaud et al 2012
Engineer &Environment
Spatio-Temporal Dynamics
Engineer ‘push’/decay ‘pull’ determine landscape environmental state & heterogeneity
Decay can stabilize engineer populations
Decay changes environmental expectations based on feedbacks
2. Dynamics of engineer consequence
Wright, J. P. 2009. Linking populations to landscapes: Richness scenarios resultingfrom changes in the dynamics of an ecosystem engineer. Ecology, 90: 3418–3429
EngineerEnvironmentSpecies richness
Outline
Ecosystem Context
Cause-Effect, Time & Space
Identity & Ecosystem Functioning
Frontiers Beyond & Within Ecology
Engineer(s) > Structural state(s) > Abiotic state(s) > Biotic state(s) > Ecosystem function(s)
A species
A ‘relatively uniform’ consortium of species
A divergent assemblage w. some net effect
Outline
Ecosystem Context
Cause-Effect, Time & Space
Identity & Ecosystem Functioning
Frontiers Beyond & Within Ecology
Ecosystem Engineering & …
Management
Evolution
Biogeochemistry
Geomorphology
Trophic Interactions
Ecosystem Engineering & Management
Humans as ecosystem engineers ‘par excellence’ (Jones et al. 1994) –Largely ignored! Why?
Lessons from nature’s engineers for human engineers?Sustainability; flexibility, adaptability & resilience; …
Ecosystem engineers, restoration, conservation, environmental management – Manage species that manage environments!Byers et al. 2006; beaver for wetlands; cows on ski slopes; vegetation on historic ruins; sheep, grass & dykes, forests & avalanches; …
Ecosystem Engineering & Evolution
Paleo-engineering (Erwin 2008)Benthic bulldozers, stout razor clams, sticklebacks
Adaptation to vs. of the abiotic environmentExtended phenotype, third helix, niche construction
Eco-Evo dynamics – Same time framesFrogs in beaver ponds, exotic Caulerpa & native bivalves
How ecosystem engineering might be usefulAbiota do not evolve they develop in response to biota & have a ‘life’ of their ownEngineering process & consequence usefully distinguished these
Ecosystem Engineering & Biogeochemistry
Abiotic controls on process rates
Engineers alter those controls in space & time
This can be integrated within & between ecosystems, but rarely is. Why?
Gutiérrez & Jones 2006
Ecosystem Engineering & (Bio-)Geomorphology
A lot of ecosystem engineering occurs through a geomorphic interface
(Bio-)geomorphology is more informed by geomorphology than ecology (ecosystem engineering) & ecosystem engineering is more informed by ecology than (bio-)geomorphology
Both disciplines have different knowledge & skill sets (concepts, methods, models, …)
The reciprocal dynamic (ecology < > geomorphology) is an emerging frontier
Jones 2012
Ecosystem Engineering & Trophic Interactions
All engineers ‘eat’ & get eaten
Engineering can affect trophic interactions
The two processes interactKelp forests; Bay of Fundy diatoms, amphipods & sandpipers; crayfish & mayflies; ants in the Negev, …
How best to integrate (Kefi et al. 2012)?Food web network structure & dynamics, energy & nutrient flow?
"Sure, kid. You start by working for the ecosystem, but pretty soon you figure out how to get the ecosystem
working for you!"