#11 Succession, trajectories,

and references
Sapaburg Urwald, Germany
 Change is a fundamental
characteristics of ecosystems
• Physical environment
– Climate changes
– Ice, volcano, earth cracks
– Soil development
• Community structure
– Temporal development and change in ecosystem
structure and function (pioneers →late successional
species)
• Genetic constitution of organisms
– Mutations and natural selection ( → evolution)
 Succession
• Sequence of plant, animal and microbial
communities that successively occupy an area
over time
• Process of change by which biotic
communities replace each other and by which
the physical environment change over time
• Sere is the consecutive communities (seral
stages)
 Primary succession
Colonization and establishment of species on land,
which has not previously been vegetated or where
plant and soil have been removed.
• Rocks
• Dunes and drifting sand
• Erosion
• Vulcano
• Glacier
• Human disturbed areas, e.g. after mining,
construction work
No biotic legacy
 Examples - primary succession
• Dune formation → lichens → grasses →
bushes → trees/forest
• Glacier melting → lichens/moss → herb/grass
→ bushes → trees/forest
• Lake → sedges → sphagnum → /dwarf shrubs
→ trees/forest
 Primary succession - Characteristics
• No biota present. They have to colonize the area. Often
wind dispersed and stress tolerant species
• Soil development, accumulation of organic matter in
soil and biomass
• Nitrogen accumulation, biotic fixation
• Microclimate often extreme and no protection from
vegetation
• The rate of change is depending on parent material and
climate
• Slowly development of biodiversity and the structural
complexity
 Secondary succession
Regrowth and recolonization on land previously
vegetated.
Reestablishment of the species, structures and
functions of existing ecosystems after
disturbances

Biotic legacy
Secondary succession – characteristics
First stage:
• Seeds, seedlings, or stools are present, enables fast
recovery after disturbance
• Light demanding species, seeds are triggered by light
or fire
• Species adapted to disturbance
Later stages
• Diversity high but may eventually decrease over time
• Biomass increases, but reach a steady-state
• Nutrient use efficiency high
• Rate of change is depending on species composition
and the degree of disturbance
 Cyclic succession

A cyclic repetition of the (entire) seral sequence

E.g.
• Heather (pioneer, building, mature,
degeneration phases)
• Primeval deciduous nemoral forest
(regeneration, early biostatic, late biostatic,
degeneration, senescence)
 What determines succession?
• Climate
• Soil/geology
• Organisms
• Time
• Disturbances
– Gaps
– Fire
– Wind
– Water
– Volcanos
– Pathogens
– Herbivorer
– Humans
Pickett et al. 2008
 Succession - Mechanisms
Autogenic - internal factors cause changes, e.g. seed
dispersal, accumulation of organic matter, changes in
water balance and nutrient availability
Allogenic – external factors causes changes in the
environment e.g. wind, fire, flooding, non-anthropogenic
climate change, grazing
Biogenic mechanisms - refer to a sudden change in the
biota, which has a major influence on succession (e.g.,
introduction of herbivores, pathogenes (Elm disease),
invasive species
Disturbances – events, which disturb or damage the
ecosystem, the populations or change the availability of
resources
 Climax communities
Plant community that no longer undergoes changes in species
composition due to succession – a hypothetic end stage
where the climax community is maintained.
• Mono-climax – This states that every region has only one
climax community, toward which all communities are
evolving (Clements 1916)
• Poly-climax - There are many different climax communities
that can be recognized in a particular region (Tansley 1939).
These climaxes are controlled by soil moisture, parent
material, activity of animals, topography, and other factors.
Succession develops towards different communities – a
mosaic of habitats and disturbed areas
• Climax forest – a self-replacing community that persists
over several generations. Gap-dynamic, micro-succession,
e.g. Suserup forest
Trajectories
 Trajectory
• An ecological trajectory is one that describes
the developmental pathway of an ecosystem
through time.
• The physical environment of the restored
ecosystem is capable of sustaining
reproducing populations of the species
necessary for its continued stability or
development along the desired trajectory.

SER Primer, 2004

 What is the difference between the
concepts of succession and trajectory?
A summer day in Denmark 11.000 years ago
Denmark 6000 years ago ……..and 2000 years ago
The Danish forests anno 2020
Untouched forest 2012, Suserup, Denmark
Palaeoecological information
 But what about the nature content we
observe today?

Quercus robur Melampyrum nemorosum

Boloria euphrosyne
Lychnis flos-cuculi Crex crex (CC BY 2.0)
Habitat share in Gribskov through time
Gribskov, Denmark.
Grib Skov, Denmark trajectory

Overballe et al., 2013

 Naturnationalpark Gribskov -
Ecological restoration?

• 1300 ha
• Release of cattle, red deer and moose, in addition to fallow deer, row
deer and sika deer already present
• Blocking of ditches
• Fellings to increase structural diversity
• Veteranisation (intentional damaging) of trees

17/09/2021 31
 The forest history
Hazel
in Denmark
forest birch Scots pine elm lime beech
epoke

poplar oak alder ash spruce

beech

Young lime

Old lime

Hazel
Birch pine
 The reference landscape (pre-neolithic)
Wood pasture hypothesis High forest hypothesis
Eriksberg, Halland, Sweden, 2012
Dear-garden, Copenhagen, Denmark
Lüneburger Heide, Germany
Oxbøl, Denmark
 Mammal keystone species
Extinct Extinct
Denmark BP Europe
Aurochs Bos primigenius Urokse 2500 1627
Bison / visent Bison bonasus Bison 2000 1911
Moose Alces alces Elg 4500. To be
reintroduced
Wild horse Equus ferus Vildhest 4500
Red dear Cervus elaphus Krondyr ++++
Fallow dear Dama dama Dådyr Reintroduced
Row dear Capreolus capreolus Rådyr +++++
Wild boar Sus scrofa Vildsvin 1801
Beaver Castor fiber Bæver 2000,
reintroduced
Brown bear Ursus arctos Bjørn 4800
Wolf Canis lupus Ulv 1813/1899
Fox Vulpes vulpes Ræv +++++
Lynx Lynx lynx Los 5000
 Class discussion
Wood pasture hypothesis High forest hypothesis
 Open-ended restoration
Based on Hughes et al. 2012 and previous class
discussions, please answer the following:

a) What characterises an open-ended restoration?

b) What situation and factors (discussed in class and
others) suggests an open-ended approach?
c) Is open-ended restoration more relevant in certain
parts of the world? Where?
d) Is an open-ended approach relevant in other
ecosystem management contexts than restoration?
E.g. rewilding, rehabilitation, ecological engineering?
 Bradshaw et al.
Based on Bradshaw et al. (2003) and previous class discussions,
please answer the following:

a) Does a single set of equilibrium baseline conditions exist?

b) Summarise arguments for and against the wood pasture
hypothesis.
c) Summarise arguments for and against the high forest
hypothesis.
d) What is the scientific evidence of these arguments?
e) Is the debate relevant in other ecosystem management
contexts than restoration? E.g. rewilding, rehabilitation,
ecological engineering?
 Vera et al.
Based on Vera et al. (2006) and previous class discussions,
please answer the following:

a) What does oak and hazel presence indicate?

b) Summarise arguments for and against the high forest
hypothesis.
c) Summarise arguments for and against the wood
pasture hypothesis.
d) What is the scientific evidence of these arguments?
e) Is the debate relevant in other ecosystem
management contexts than restoration? E.g.
rewilding, rehabilitation, ecological engineering?
 Svenning
Based on Svenning et al. (2002) and previous class
discussions, please answer the following:

a) What is the link between pollen records and

vegetation structure?
b) What is suggested about the geographical spread of
high forest and more light open habitats?
c) List drivers of light open habitats?
d) What is the scientific evidence of these arguments?
e) Is the debate relevant in other ecosystem
management contexts than restoration? E.g.
rewilding, rehabilitation, ecological engineering?
 What is open-ended restoration?
• Minimal intervention (by humans)
• Acceptance of future trajectories (ecological change)
• Habitats formed by autogenic processes (regeneration,
succession) and allogenic processes (e.g. plant
dispersal, river flooding)
• Colonization and extinction are allowed
• Climate change influence the trajectory
• Disturbance at different scales (in space and time)
• Change may be stepwise, sudden and irreversible or
more gradual.
Hughes et al., 2012
 When and where is open-ended
restoration relevant to consider?
• In remote areas
• Large scale projects (large areas)
• In systems to be affected by climate change (temperature, not least
precipitation)
• In systems that are naturally very dynamic
• Where starting points are novel
– After mining, agriculture,
– Meaningless to go back to reference level
– Invasive species can’t be managed
• When ongoing management is very expensive
• Give nature back to nature
• Not: abandon and ignore. Rather: Monitor and learn.
• Where else?

Hughes et al., 2012

 Wood pasture development
sensu Vera (2012)
• Trees colonized after last glaciation
• Landscape was kept semi-open by large herbivores
(aurochs, visent, tarpan etc)
• Man cut the trees and cultivated the land
• Trees and animals lost their home land
(Red dear, Elk, European bison (visent), Wolf, Lynx, Aurochs,
Tarpan)
• Pastures were established and other organisms
benefited
• This semi or half-natural landscapes were very
biodiverse.
 Wood pasture hypothesis
Assets:
• Explain the early Holocene landscape
– Quercus, Pinus, Corylus
– Large mammals
Weakness:
• Lack of palaeoecological evidence
• Open structures rather relates to fire than
ungulates
• Lack of non-intervention modern analogues.

Suggested by Frans Vera, 2000

Suserup Denmark
Suserup Denmark
Høstermark Skov
Denmark
 High Forest hypothesis
Assets:
• Consistant with palaeoecological data (pollen,
fossils, insects)
• Modern analogues in non-intervention forests

Weaknesses:
• Where were all the light demanding species
(how had they surrived)?
 Closed (high) Forest Dominate
Normal uplands (well drained loamy soils)

Less dense forest were present on

• Floodplains (histo and gleysols)
• Infertile soils (sandy podzols)
• Chalklands (dry rendzinas)

Svenning 2002
 High forest or wood pasture?

Climax (potential natural vegetation) is a construction without

taking into consideration any impact of indigenous large
harbivores (aurochs and tarpans). Vera, 2013
 Two different goals
1. Open-ended trajectory restoration
– Processes are free
– Stochastic development
2. Target oriented restoration
– Specific goals
– Species conservation
– Natura2000