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DynamicDynamic andand SuccessionSuccession ofof EcosystemsEcosystems

Kristin Heinz, Anja Nitzsche Basics of Analysis 10.05.06 Structure • Ecosystem dynamics –Basics –Rhythms – Fundamental model

• Ecosystem succession –Basics – –Diversity – Examples Ecosystem dynamics

• Because of the open character there is a flow between different • It is a flow of – Energy → transformation –Stocks – Dynamics in populations –… Ecosystem dynamics

• Biological rhythms – Circardian rhythm – Annual rhythm – Longer annual rhythm –Tides – Lunar rhythm –… Biological rhythms

• Circardian rhythm – Production of – Vertical drift of limnic and marine – Roost fly of birds in winter

• Annual rhythm – Falling down of the leaves – Hibernation – Bird migration – Diapause Ecosystem dynamics

• Longer annual rhythm – In

• Tides – Along the cost side very different and complex structure of time – Characteristic vertical zoning of the animals and alga Ecosystem dynamics

• Lunar rhythm – Agitation in migration by birds, tropical mammals and insects Ecosystem dynamics

• Human made rhythms – Land use activities – Change of land use – Emission dynamics – Environmental policy – Global change – Continuous climate change –… Fundamental model of ecosystem dynamics

Stored 4 Renewal Conservation 2 - Accessible Carbon, - k-Strategy - Nutrients ad Energy - Climax Ca - Consolidation pital (Storage)

LY W R O (Miineralisation) A L (Adult Stage) P S ID L Y

1 Exploitation Creative 3 - r-Strategy Destruction - Pioneers -Fire - Opportunists -Storm -Pest - Senescence (Juvenile Stage) ( Incorporation) Organization Connectedness

(Holling 1986) A hypothetic trajectory of the adaptive cycle

Maturity / Conservation

Renewal / Exergy Reorganization stored Release / Creative destruction

Pioneer stage / Exploitation

connectedness Disturbance

• Stability – Ability of an ecosystem to recover or to return to the original constitution after disturbance • Resiliency – Dimension for the ability of an ecosystem to survive a disturbance • Capacity – Intensity of resiliency factors which can be buffer in an ecosystem – Only stable ecosystems can buffer Disturbance

• Difference from the original ecological factors • Direct and indirect damages Ecosystem Succession - Definition

• A fundamental concept in

• Refers to more-or-less predictable and orderly changes in the composition or structure of an ecological

• Initiation by:

¾ Formation of a new, unoccupied (e.g., a lava flow or a severe landslide) → primary succession

¾ Some form of disturbance (e.g. fire, severe windthrow) of an existing community → secondary succession Ecosystem Succession - Definition

• The trajectory of ecological change can be influenced by:

¾ site conditions

¾ the interactions of the species present

¾ more stochastic factors such as availability of colonists or seeds Ecosystem Succession - Definition

• Stable end-stage called climax, sometimes referred to as the 'potential ' of a site, shaped primarily by the local climate

• Has been largely abandoned by modern ecologists in favour of nonequilibrium ideas of how ecosystems function

• Most natural ecosystems experience disturbance at a rate that makes a "climax" community unattainable

Æ climate change Æ expansions and introductions Ecosystem Succession - Factors

• Succession usually occurs in areas where no other species offer in the area

• The type of ' that occupy areas in such circumstances depends on a number of factors

• Climate → Temperature, precipitation, sunlight

→ pH, composition of the soil

• Human Intervention → pollution, acid rain Ecosystem Succession – Energy Flow

• Energy flow is fundamentally changed → demonstrated in the quantity of standing crop in the ecosystem

• During early seral stages → energy inputs > outputs

• Disturbance by external factors → the energy loss > inputs

• Accumulation of energy as → high metabolic rates → high which maximises the energy flow in the system Ecosystem Succession - Productivity

• Increases proportional to the amount of standing crop

• The percentage gross productivity is fixed as net productivity is not continuous with progression

• Early seral stages: small short-living plants dominate, have a high yield and individual plants require very little energy for maintenance Æ r-selected life-histories

• Later seral stages: large long-living plants dominate which use high levels of their gross productivity for respiration to maintain their bodies Æ k-selected life-histories Ecosystem Succession - Diversity

• Number of species progresses rapidly as plants and animals colonise the area

• In later seres the rate of increase decreases

• Increasing interspecific competition → it is the intermediate seres which contain the largest number of species present at any one time during the succession Ecosystem Succession - Trophic Structure

• Early seres are short, linear food chains which are easily upset if one element in the is removed

• As the succession progresses the ecosystem becomes more layered and increases creating a complex

• The more complex food webs → greater stability → allows alternative energy flows when one element of the food chain is disrupted Ecosystem Succession - 1st Example

• Primary Succession: colonisation of bare rock

¾ Pioneer community: lichens → provide enough nutrients to support a community of small plants such as mosses → typically replaced by ferns

¾ With erosion of rock and increasing amounts of organic material a large layer of soil is gradually built up

¾ This soil allows plants such as grasses and small flowering plants to grow followed by shrubs and → climax community? Ecosystem Succession - 2nd Example

• Secondary succession: after forest fire

¾ Spores, seeds and vegetative organs may remain viable in the soil

¾ Influx of animals and plants through dispersal and migration from the surrounding area

¾ Succession does not begin with pioneer species but with species from intermediate seres