Inputs Outputs Ecosystem Production Consumption Decomposition Element cycling FEE so far Energy flow Chemical transformation The extended physics of biota.

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!"