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

3. 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

4. http://www.outdoorescapesnewhampshire.com/Photos/BeaverDam.jpg http://ci.marysville.wa.us/publicworks/swm/docs/beaver_files/Beaver%20Pic.jpg Beaver → Dams → Hydrology, sediments → Biogeochemistry, habitat Two of countless examples

5. http://library.thinkquest.org/05aug/00183/How%20Hurricanes%20Impact%20the%20Reef.htm Coral reefs → Wave attenuation → Refugia

6. Xxxxx →Xxxxxx → Xxxxxx → Xxxxxxx

7. Outline  Ecosystem Context  Cause-Effect, Time & Space  Identity & Ecosystem Functioning  Frontiers Beyond & Within Ecology

8. A place with all the living & non-living interacting – Tansley 1935 Abiotic > Abiotic: Physical & chemical processes e.g., erosion & deposition (material vectoring), … ; redox, precipitation, hydrolysis, … Biotic > Biotic: Direct energy, material, information transfer e.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) – Trophic e.g., plant nutrient uptake & OM production, urine, feces, microbial mineralization Physical ecosystem engineering process – Non-trophic e.g., beaver dam & pond, root macropore & drainage, coral reef wave attenuation … All Interacting e.g., nutrient cycling; direct abiotic resource competition; physical ecosystem engineering consequence (biogeochemical heterogeneity, habitat, engineer feedbacks,…); …

9. 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 & Dissimilation Physical Ecosystem Engineering Abiotic Determinism BIOTA Direct Energy, Material, & Information Transfers

10. 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

11. Outline  Ecosystem Context  Cause-Effect, Time & Space Definitions Framework: Mostly Temporal Space Spatio-Temporal Dynamics  Identity & Ecosystem Functioning  Frontiers Beyond & Within Ecology

12. Jones & Gutiérrez 2007 Definitions Physical Non-Trophic Abiotic resources Abiotic conditions Biotic effect Process Consequence Ecosystem

13. Jones et al. 2010 Framework: Mostly Temporal Process, consequences, system dynamics Magnitudes

14. Engineering Process Engineer > structural change > abiotic change Exemplify!

15. 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?

16. Structural decay (D) 1. Agents? 2. Why occur? 3. Determinants? 4. Offset? 5. Legacy fate? 6. Measure? Units?

17. Net structural change (F-D) 1. Why relevant? 2. Measure? Units?

18. Abiotic change 1. Why occur? 2. Determinants? 3. Measure? Units? 4. Relationship between abiotic change & structural decay?

19. Engineering Consequence Structural/abiotic change > Biotic change & engineer feedbacks Exemplify!

20. 1. Why occur? 2. How predict? Expectations? 3. Measure? Units? 4. Why feedbacks relevant? 5. Kinds of feedback? Time scales? Consequences?

21. Space Engineer Engineering requirements Structural- & abiotic-state dependence Other species Engineered habitat specialists Unmodified habitat specialists Habitat generalists Energetic & material connectance Engineered Unmodified EMO

22. 1. General model of environmental dynamics Raynaud et al 2012 Engineer & Environment Spatio-Temporal Dynamics

23. Engineer ‘push’/decay ‘pull’ determine landscape environmental state & heterogeneity Decay can stabilize engineer populations Decay changes environmental expectations based on feedbacks

24. 2. Dynamics of engineer consequence Wright, J. P. 2009. Linking populations to landscapes: Richness scenarios resulting from changes in the dynamics of an ecosystem engineer. Ecology, 90: 3418–3429 Engineer Environment Species richness

25. Outline  Ecosystem Context  Cause-Effect, Time & Space  Identity & Ecosystem Functioning  Frontiers Beyond & Within Ecology

26. 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

28. Outline  Ecosystem Context  Cause-Effect, Time & Space  Identity & Ecosystem Functioning  Frontiers Beyond & Within Ecology

29. Ecosystem Engineering & … Management Evolution Biogeochemistry Geomorphology Trophic Interactions

30. 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; …

31. Ecosystem Engineering & Evolution Paleo-engineering (Erwin 2008) Benthic bulldozers, stout razor clams, sticklebacks Adaptation to vs. of the abiotic environment Extended phenotype, third helix, niche construction Eco-Evo dynamics – Same time frames Frogs in beaver ponds, exotic Caulerpa & native bivalves How ecosystem engineering might be useful Abiota do not evolve they develop in response to biota & have a ‘life’ of their own Engineering process & consequence usefully distinguished these

32. 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

33. 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

34. Ecosystem Engineering & Trophic Interactions All engineers ‘eat’ & get eaten Engineering can affect trophic interactions The two processes interact Kelp 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?

35. "Sure, kid. You start by working for the ecosystem, but pretty soon you figure out how to get the ecosystem working for you!"