Course2.1.1: Basics of Ecosystem Analysis

May 24, 2006 Ecosystem ProtectionConcepts

A Ecosystem ProtectionII. à W. Windhorst -Ecosystem Integrity, theConcept à Pres. Timo -Ecosystem Integrity, Applications à Pres. Timo

TheUNCBD Ecosystem Approach –CaseStudies à W. Windhorst -ForestEcosystems à Pres. Pawel

Ecosystem Theory. à F. Müller B OrganizingSalzau -NetworkTheory à Pres. Kamil Web pages -Thermodynamics à Pres. Aiko Transfer -HierarchyTheory à Pres. Pawel Workingplans -Gradient Principles à Pres. Lech Problems August Macke: Felsige Landschaft, 1914, Aquarell, 24 ×20 cm, Land: Deutschland, Stil: Expressionismus. What are theories? Popper: Abstract "Theory is the fishing net description, that scientists cast explanation and to catch the world, organization to explain it of a scientific and to control it." disciplin.

Theories are - sets of hypotheses empirically or deductively - non-evaluative based, aggregating and - capable of being integrating falsified representations - prognostic of the proved - applicable to knowledge of a single cases scientific disciplin. - directing the development, What orientation, and are Theories structure of the disciplin theories? Which theories are relevant for ecosystem compre- hension? Approaches: Farina: - Hierarchy Theory "Theories are interpreting - Fractal Geometry the complexity and - Chaos Theory heterogeneity of the - Catastrophe environment (landscape)" Theory - Self-Organization Naveh & Liebermann: - Information Theory "The organization of - Thermodynamics complexity in landscape - Theory of Which structure and function Dissipative theories has become a central Structures issue in contemporary - Percolation Theory are landscape ecology." - Metapopulation relevant Theory for - Systems Source- ecosystem Landscape Sink - Spatial Pattern compre- Theories Analysis hension? Cybernetics Network Theory Regulation and Control Flow Analysis in in Complex Systems, Complex, Interacting Feedback Loops, Systems Homoeostasis, e.g. Electrical Circuits Optimization

Ecological Modelling Trophic Stability Theory Flows, Which Conceptual Landscape Chains, and Networks theories and Ecosystem Models Food Webs Indirect Effects are relevant for ecosystem Ecosystem compre- hension? Theories Information Theory Hierarchy Theory Improbability of the Analysis of Subsystems Occurence of an Event with Asymmetric Inter- actions Syntactic, Semantic, Scales, Holons, Pragmatic Information Constraints and Filters

Measures of Diversity Principles and Methods Measures of Complexity of Scale Distinction Which Ascendency theories - Total Throughflow Fundamentals for the are - Flow Articulation Existence of Emergence relevant for ecosystem A = I * T compre- Ecosystem hension? Theories HolonLevel (+1) SpatialExtent: High Frequencies: Low

MinorInteractions Multiple Interactions HolonLevel (0) Filter

Constraints

Holon Level SpatialExtent: Small (-1) Frequencies: High

Hierarchiesand theproblemsof scale Synergetics Thermodynamics Detecting the Principles Analyzing Balances and of Self-Organization in Transfers of Energy Physical, Chemical, and Corresponding Biological and Human Principles Systems

Self-Organization Energetic Principles of Ecosystems and of Ecological Landscapes, Self-Organization Gradients and Dissipative Structures Which Emergent Exergy, Entropy, Properties Emergy theories are Orientors relevant Principles of Undisturbed for Ecosystem Development of ecosystem Ecosystems and Theories Landscapes compre- hension?

(Voluntary) Homework: Find examplesfor…

terrestrial aquatic

-circadianerhythms o o

-lunearperiodics o o

-tideperiodics o o

-seasonalrhythms o o

-longtermrhythms o o (modifiedafter Holling1995; Hollingetal.2000)

AVAILABLE NUTRIENTS

Decomposition EXE R G Y C ON S U M P TION K-strategy climax

pest fire storm toanother attractor r-strategy pioneer Lotsof dead biomass opportunist

Storedexergy Holling’s conceptual model incorporates both Clementsian linear succession and Gleasonian independent, species-level disordered behaviour, which are integrated into a complexity based framework with insights gleaned from catastrophe theory, chaos theory and self-organization theory.

FigurefromG. Zurlini Whatisa succession?

.... an orderedprocessof communitydevelopment includingregularchangesof thespeciescompositions

.... changesoccuringdueto variationsof thespeciesinteractions

.... changesoccuringdueto internalprocesseswithinthe community

.... changesin communitystructureeffectingchangesin the physicalecosystemfunctions -energy -whichdirection? -water -whichdirection? -nutrients -whichdirection? Typesof successions:

-autogeneoussuccessions ---> consequencesof internalprocesses

-allogeneoussuccession ---> consequencesof externalinfluences

-primarysuccession ---> „natural“development

-secondarysuccession ---> followingprimarysuccessionswhichhave developedunderman‘s disturbing influences Typesof successions:

-progressive successions ---> complexification

-retrogressivesuccession ---> consequencesof disturbances simplification Climax:

-final and „stable“communityof a developmentalsequence

-climaticalclimax theoreticalcommunitywhichisthetypicalresult of successionswithina certain(climatic) area

-edaphicclimax resultof succession, effectingmodificationsof the climaticclimaxcommunitydueto physical modificationsof thesites Threeexamples forecosystem development fromODUM

a. Microorganisms in a haysolution

Aus: H.J. Müller (1984) Threeexamples forecosystem development fromODUM

a. Microorganisms in a haysolution

b. Plant successions afterfireand in dune systems

Aus: H.J. Müller (1984) Threeexamples forecosystem development fromODUM

a. Microorganisms in a haysolution

b. Plant successions afterfireand in dune systems

c. Primaryproduction and respiration throughout successions

Aus: H.J. Müller (1984) A mutualhypothesis fromWeber et al. (1989) A mutual hypothesisof ecosystemdevelopment in phasesof biologicalprocessdominance

Import

+ Autocatalytic system + Internal New Flows, Cycles element Export Integration of new elements (species) into an autocatalytic system will take place, if loss is decreased, internal cycling is optimized, and dampening capacity increases (Weber et al. 1989) Efficiency of the whole as a parameter of selection

weber.pre Maximum Size due to the Constraints, Provided by the Site Conditions

"Resulting State" "Goal" "Attractor"

Time / Maturity / Stage of Development Maximum Size due to the Constraints, Provided by the Site Conditions

Dynamics of a potential higher hierarchical level

Dynamics of a potential lower hierarchical level

Time / Maturity / Stage of Development Ecophysiological and functional orientors System System -lossreduction dynamical organizational -storages -biomass orientors orientors -biomass/production ratio -buffercapacity? -complexity -harvest/throughput -stability? -hierarchicallevels ratio -(meat)stabilty? -semiautonomy -transpiration -resilience? -feedbackcontrol -respiration -resistance? -holistic -respiration/biomass -signalfiltering? determination ratio -criticallity -resourceutilization -connectedness -utility -overconnectedness -intraorganismic -redundancy storages -metabolicquotients Do ecosystemsalwaysfollowtheorientors? Do ecosystemsalwaysfollowtheorientors?

No. 4 Renewal Conservation 2 - Accessible Carbon, - k-Strategy - Nutrients and Energy - Climax - Consolidation

(Miineralisation) (Adult Stage) RADIPLY SLOWLY

schnell

S t or e d C a p i l ( g e) 1 Exploitation Creative 3 - r-Strategy Destruction - Pioneers - Fire - Opportunists - Storm - Pest - Senescence (Juvenile Stage) ( Disturbance Incorporation) Organization

4box1.pre Connectedness

Catalogueof questions

•Whatisa successionand howcanwedistinguishsuccessiontypes? •Find examplesforprogressive and retrogresivesuccessions. •Whicharethetypicaldevelopmentalphasesof lakecommunities •on an annualbasis? •within10 years? •within1000 years? •Whatisan orientor? •Whicharethedifferencesbetweentheconceptsof stability, resilience, elasticity, and buffercapacity? •Whicharethecomponentsof the‚adaptive cycle‘referringto •a bacteriapopulation •thedune systemof theislandof Sylt? •theman-environmentalsystemof Schleswig-Holstein duringthepast250 years?

StructurizedSystems State withConcentrationGradients StructurizedSystems State withConcentrationGradients

Diffusion, Dispersion, Dissipation

ThermodynamicEquilibrium (no Gradients) StructurizedSystems State withConcentrationGradients

Diffusion, Dispersion, Dissipation

ThermodynamicEquilibrium (no Gradients)

Dissipative Self-Organisation

StructurizedSystems State withNew Gradients Self-Organization:

Spontaneous Creation of Macroskopic Structuresfrom Microskopic Disorder

Gradient Formation Principlesof Self-Organization: Principlesof Self-Organization:

Exergy Import

Exergy Degradation

EntropyExport Principlesof Self-Organization:

Exergy Import ConvertibleEnergy e.g. Radiation

CO2-Input

Exergy Degradation EnergyTransformation e.g. PhysiologicalProcesses Growth Respiration

EntropyExport EnergyOutput e.g. Heat

CO2-Output Principlesof Self-Organization:

Exergy Import ConvertibleEnergy e.g. Radiation

CO2-Input

Exergy Degradation EnergyTransformation e.g. PhysiologicalProcesses Growth Respiration

EntropyExport EnergyOutput e.g. Heat

CO2-Output

Gradient Formation Principlesof Self-Organization:

Exergy Import Systems State Far fromEquilibrium

InternalControl/ Regulation

SymmetryBreakingOrganization Exergy Degradation Cooperativityof Subsystems ThermodynamicOpenessof System Constraintsin Hierarchies Meta StabilityafterSmall Impulses EntropyExport Fluctuationsin Phase Transitions Historicityand Irreversibility

Gradient Formation

CriteriaafterEbeling(1989) System in a Normal State System in an ExcitedState

System in a Normal State Exergy Gradient En e rgy Input Energy Potentially U t i li z a ble Captured CaseA: IsolatedSystem

AvailableEnergy cannotbeCaptured

Exergy Entropy Gradient Produced En e rgy En e rgy Input Energy Outp ut U til i z a b le Potentially Energy - U t i li z a ble Captured Emitted Non CaseB: IntegratedSystem

AvailableEnergycan beCapturedand Utilized

Exergy Degrading Network En e rgy Input Outp ut U t i li z a ble Gradients and Non-Equilibrium Self-Organization in Principle Leads to Complexifying

SCHNEIDER & KAY (1994): Dynamics "If a system is moved away from with thermodynamic equilibrium, by the application of a flow of exergy, an it will utilize all avenues available, Increasing that is build up as much dissipative Number structure as possible, to reduce the effects of the applied exergy gradient." of Living systems are degrading and utilizing Interrelated applied gradients by the Gradients self - organized formation of a hierarchy of nested internal gradients. Thetentative fourthlawof thermodynamics

S.E. Joergensen: Ifa systemsreceivesa through-flowof exergy, thesystem will utilisethisexergy to moveawayfromthermodynamic equilibrium. Ifthesystemisofferedmorethanonepathwayto move awayfromthermodynamicequilibrium, theoneyieldingmost storedexergy, i.e. withthemostorderedstructureor thelongestdistance fromequilibriumbytheprevailing conditions, will havea propensityto beselected.

----> optimisationof exergy storage

-biomass -organicmatter -structureand information Maturity: Optimum defined by the site's constraints

GRAD

EX (T) EX (S)

GRAD = Diversity of gradients EX (T) = Total amount of exergy captured EX (M) EX (S) = Exergy invested into the creation of the gradient structure (storage) EX (M) = Exergy necessary for the maintenance of the gradient structure (degradation) Pioneer state --- Developmental time --- Adult state ecotheo2.pre Maximum Sizedueto theConstraints, ProvidedbytheSite Conditions

"Resulting State" "Goal" "Attractor"

Time / Maturity/ Stageof Development Maximum Sizedueto theConstraints, ProvidedbytheSite Conditions

Disturbance - natural externalinputs (e.g. fire, storm, pests) - human externalinputs (e.g. clearcut, pollution,...) - senescenceof a system (e.g. one-agedforests) - dominanceof species (e.g. mossesin fens, pests) - internalfeedbackloops (e.g. socaccumulation)

Time / Maturity/ Stageof Development Maximum Size due to the Constraints, Provided by the Site Conditions

Disturbance

System A

System B

Time / Maturity / Stage of Development Maximum Size due to the Constraints, Provided by the Site Conditions

Dynamics of a potential higher hierarchical level

Dynamics of a potential lower hierarchical level

Time / Maturity / Stage of Development Structural and Network community theoretical orientors orientors Information theoretical - development of - indirect effects - cycling index orientors symbioses - life span - complexity of cycles - body size - chain length - information - r/k selection - average trophic - heterogeneity - fecundity levels - species richness - biomass - residence times - Shannon index - network - ascendency homogenization

- system throughput - network - niche diversity - system overhead amplification - specialization - connectivity - network synergism - prey specificity - utility - trophic efficiencies Thermodynamic Thermodynamic Thermodynamic orientors: orientors: orientors: exergy entropy emery and power

- exergy capture - entropy - emergy - exergy flows - minimum excess - power - exergy consumption entropy - flow activity - exergy degradation - total entropy - flux density - exergy storage production - specific exergy - specific entropy - structural exergy production Thermodynamic - dissipation orientors: gradients Thermodynamic - structure orientors: - information - gradient emergence distance from - heterogeneity - gradient degradation equilibrium - complexity Ecophysiological and functional orientors System System - loss reduction dynamical organizational - storages orientors - biomass orientors - biomass/production ratio - buffer capacity ? - complexity - harvest/throughput - stability ? - hierarchical levels ratio - (meat)stabilty ? - semi autonomy - transpiration - resilience ? - feedback control - respiration - resistance ? - holistic - respiration/biomass - signal filtering ? determination ratio - criticallity - resource utilization - connectedness - utility - overconnectedness - intraorganismic - redundancy storages - metabolic quotients Catalogueof questions:

- Are ecosystemsdissipative systems? -Are ecosystemsself-organizedsystems? Tryto check theprinciples of self-organizationafterEbeling(1989)

Tryto find good definitionsof threethermodynamicvariables: - Whatistheexergy of a system/an ecosystem? - Whatistheentropyproductionof a system/an ecosystem? - Whatisemergyand wherecanyoufind itin ecosystems?

Tryto applythethermodynamicprinciplesbycomparisonof the energybudgetsof (a) a desertand (b) a tropicalrainforestwith referenceto - exergy capture - exergy storage - exergy degradation - entropyproduction