Ecological succession

Succession after disturbance: a boreal forest one year (left) and two years
(right) after a wildfire.
Ecological succession is the process of change in the species structure
of an ecological community over time. The time scale can be decades (for
example, after a wildfire), or even millions of years after a mass extinction.
The community begins with relatively few pioneering plants and
animals and develops through increasing complexity until it becomes stable
or self-perpetuating as a climax community. The "engine" of succession,
the cause of ecosystem change, is the impact of established organisms
upon their own environments. A consequence of living is the sometimes
subtle and sometimes overt alteration of one's own environment.
It is a phenomenon or process by which an ecological
community undergoes more or less orderly and predictable changes
following a disturbance or the initial colonization of a new habitat.
Succession may be initiated either by formation of new, unoccupied habitat,
such as from a lava flow or a severe landslide, or by some form
of disturbance of a community, such as from a fire, severe windthrow,
or logging. Succession that begins in new habitats, uninfluenced by pre-
existing communities is called primary succession, whereas succession
that follows disruption of a pre-existing community is called secondary
succession. Succession was among the first theories advanced in ecology.
Ecological succession was first documented in the Indiana Dunes of
Northwest Indiana and remains at the core of ecological science
History
Precursors of the idea of ecological succession go back to the beginning of
the 19th century. The French naturalist Adolphe Dureau de la Malle was
the first to make use of the word succession concerning the vegetation
development after forest clear-cutting. In 1859 Henry David Thoreau wrote
an address called "The Succession of Forest Trees" in which he described
succession in an oak-pine forest. "It has long been known to observers that
squirrels bury nuts in the ground, but I am not aware that any one has thus
accounted for the regular succession of forests."The Austrian
botanist Anton Kerner published a study about the succession of plants in
the Danube river basin in 1863.
H. C. Cowles

The Indiana Dunes on Lake Michigan, which stimulated Cowles'

development of his theories of ecological succession
Henry Chandler Cowles, at the University of Chicago, developed a more
formal concept of succession. Inspired by studies of Danish dunes
by Eugen Warming, Cowles studied vegetation development on sand
dunes on the shores of Lake Michigan (the Indiana Dunes). He recognized
that vegetation on dunes of different ages might be interpreted as different
stages of a general trend of vegetation development on dunes (an
approach to the study of vegetation change later termed space-for-time
substitution, or chronosequence studies). He first published this work as a
paper in the Botanical Gazette in 1899 ("The ecological relations of the
vegetation of the sand dunes of Lake Michigan").[7] In this classic
publication and subsequent papers, he formulated the idea of primary
succession and the notion of a sere—a repeatable sequence of community
changes specific to particular environmental circumstances
Gleason and Clements
From about 1900 to 1960, however, understanding of succession was
dominated by the theories of Frederic Clements, a contemporary of
Cowles, who held that seres were highly predictable and deterministic and
converged on a climatically determined stable climax community regardless
of starting conditions. Clements explicitly analogized the successional
development of ecological communities with ontogenetic development of
individual organisms, and his model is often referred to as the pseudo-
organismic theory of community ecology. Clements and his followers
developed a complex taxonomy of communities and successional
pathways.
Henry Gleason offered a contrasting framework as early as the 1920s. The
Gleasonian model was more complex and much less deterministic than the
Clementsian. It differs most fundamentally from the Clementsian view in
suggesting a much greater role of chance factors and in denying the
existence of coherent, sharply bounded community types. Gleason argued
that species distributions responded individualistically to environmental
factors, and communities were best regarded as artifacts of the
juxtaposition of species distributions. Gleason's ideas, first published in
1926, were largely ignored until the late 1950s.
Modern era
A more rigorous, data-driven testing of successional models and
community theory generally began with the work of Robert
Whittaker and John Curtis in the 1950s and 1960s. Succession theory has
since become less monolithic and more complex. J. Connell and R.
Slatyer attempted a codification of successional processes by mechanism.
Among British and North American ecologists, the notion of a stable climax
vegetation has been largely abandoned, and successional processes have
come to be seen as much less deterministic, with important roles for
historical contingency and for alternate pathways in the actual development
of communities. Debates continue as to the general predictability of
successional dynamics and the relative importance of equilibrial vs. non-
equilibrial processes. Former Harvard professor F. A. Bazzaz introduced
the notion of scale into the discussion, as he considered that at local or
small area scale the processes are stochastic and patchy, but taking bigger
regional areas into consideration, certain tendencies can not be denied.[12]
Factors
The trajectory of successional change can be influenced by site conditions,
by the character of the events initiating succession (perturbations), by the
interactions of the species present, and by more stochastic factors such as
availability of colonists or seeds or weather conditions at the time of
disturbance. Some of these factors contribute to predictability of succession
dynamics; others add more probabilistic elements. Two important
perturbation factors today are human actions and climatic change.[13]
In general, communities in early succession will be dominated by fast-
growing, well-dispersed species (opportunist, fugitive, or r-selected life-
histories). As succession proceeds, these species will tend to be replaced
by more competitive (k-selected) species.
Trends in ecosystem and community properties in succession have been
suggested, but few appear to be general. For example, species
diversity almost necessarily increases during early succession as new
species arrive, but may decline in later succession as competition
eliminates opportunistic species and leads to dominance by locally superior
competitors. Net Primary Productivity, biomass, and trophic properties all
show variable patterns over succession, depending on the particular
system and site.
Ecological succession was formerly seen as having a stable end-stage
called the climax, sometimes referred to as the 'potential vegetation' of a
site, and shaped primarily by the local climate. This idea has been largely
abandoned by modern ecologists in favor of nonequilibrium ideas of
ecosystems dynamics. Most natural ecosystems experience disturbance at
a rate that makes a "climax" community unattainable. climate change often
occurs at a rate and frequency sufficient to prevent arrival at a climax state.
Additions to available species pools through range expansions
and introductions can also continually reshape communities.
The development of some ecosystem attributes, such as soil
properties and nutrient cycles, are both influenced by community
properties, and, in turn, influence further successional development. This
feed-back process may occur only over centuries or millennia. Coupled
with the stochastic nature of disturbance events and other long-term (e.g.,
climatic) changes, such dynamics make it doubtful whether the 'climax'
concept ever applies or is particularly useful in considering actual
vegetation.
Secondary succession

An example of Secondary Succession by stages:

1. A stable deciduous forest community
2. A disturbance, such as a wild fire, destroys the forest
3. The fire burns the forest to the ground
4. The fire leaves behind empty, but not destroyed, soil
5. Grasses and other herbaceous plants grow back first
6. Small bushes and trees begin to colonize the area
7. Fast growing evergreen trees develop to their fullest, while shade-
tolerant trees develop in the understory
8. The short-lived and shade intolerant evergreen trees die as the larger
deciduous trees overtop them. The ecosystem is now back to a similar
state to where it began.
Successional dynamics following severe disturbance or removal of a pre-
existing community are called secondary succession. Dynamics in
secondary succession are strongly influenced by pre-disturbance
conditions, including soil development, seed banks, remaining organic
matter, and residual living organisms. Because of residual fertility and pre-
existing organisms, community change in early stages of secondary
succession can be relatively rapid.
Secondary succession is much more commonly observed and studied than
primary succession. Particularly common types of secondary succession
include responses to natural disturbances such as fire, flood, and severe
winds, and to human-caused disturbances such as logging and agriculture.
In secondary succession, the soils and organisms need to be left unharmed
so there is a way for the new material to rebuild.[4]
As an example, in a fragmented old field habitat created in eastern Kansas,
woody plants "colonized more rapidly (per unit area) on large and
nearby patches".[16]
Secondary succession: trees are colonizing uncultivated fields and
meadows.
Secondary succession can quickly change a landscape. In the
1900s, Acadia National Park had a wildfire that destroyed much of the
landscape. Originally evergreen trees grew in the landscape. After the fire,
the area took at least a year to grow shrubs. Eventually, deciduous trees
started to grow instead of evergreens.[15]
Secondary succession has been occurring in Shenandoah National
Park following the 1995 flood of the Moorman's and Rapidan rivers, which
destroyed plant and animal life.[17
Seasonal and cyclic dynamics
Unlike secondary succession, these types of vegetation change are not
dependent on disturbance but are periodic changes arising from fluctuating
species interactions or recurring events. These models modify the climax
concept towards one of dynamic states.
Hydrarch Succession [18]

Succession initiated with establishment of the pioneer communities in a wet

area is termed hydrarch succession and the succession stages are
hydrosere and halosere.

1. Hydrosere: When succession starts in the fresh water ecosystem like

ponds, pools, lakes and marshes.
2. Halosere: When it start in saline water ecosystem, for example-
mangroves, coral reefs. [19]
Succession in the pond begins by colonization by the pioneers like the
phytoplankton and finally terminates into a forest, which is climax
community. The whole process of succession of the hydrosere is further
subdivided into a number of sub stages depending on the kind of organism
dominating a stage.
The whole process of succession of hydrosere is further subdidvided into a
number of sub-stages depending on the kind of organism dominating a
stage:
1. Phytoplankton stage: In this initial stage , the pond water is poor in
nutrients and devoid of much life. At this stage the water is incapable of
supporting larger life forms. In such situations, phytoplanktons consisting of
microscopic algaoe begin to multiply and they quickly become the pioneer
of colonizers. As the phytoplanktons and the dependent animal population
die, decomposer organisms increase in number and bring about
decomposition , which results in the release of minerals and enrichment of
aquatic habitats.
2.Submerged stage : the habitat which is now shallower and richer in
nutrients, where light is available upto a certain depth becomes suitable for
growth of rooted , submerged hydrophytes like Hydrila , Potamogeton,
Vallisneria etc. these grow at various depths mostly rooted in the muddy or
sandy bottom depending on the species and also on the clearness or
turbidity of water.
3. Floating stage: The pond is now colonized by plants which are rooted in
mud but their leaves reach water surface and float. Due to the growth,
death and decay of this organism, the water level by now become very
much decreased making the pond much shallower, evaporation of water
from the adjacent area also contribute in this process[18].
Xerarch Succession[18]
Succession initiated with establishment of the pioneer communities in drier
area is termed xerarch succession and successional stages are called
xerosere .

1. Lithosere : When succession takes place on bare rocks.

2. Psammosere :When succession takes place in a sandy area, like
sandy dunes.
Causes of plant succession[edit]
Autogenic succession can be brought by changes in the soil caused by the
organisms there. These changes include accumulation of organic matter in
litter or humic layer, alteration of soil nutrients, or change in the pH of soil
due to the plants growing there. The structure of the plants themselves can
also alter the community. For example, when larger species like trees
mature, they produce shade on to the developing forest floor that tends to
exclude light-requiring species. Shade-tolerant species will invade the area.
Allogenic succession is caused by external environmental influences and
not by the vegetation. For example, soil changes due to erosion, leaching
or the deposition of silt and clays can alter the nutrient content and water
relationships in the ecosystems. Animals also play an important role in
allogenic changes as they are pollinators, seed dispersers and herbivores.
They can also increase nutrient content of the soil in certain areas, or shift
soil about (as termites, ants, and moles do) creating patches in the habitat.
This may create regeneration sites that favor certain species.
Climatic factors may be very important, but on a much longer time-scale
than any other. Changes in temperature and rainfall patterns will promote
changes in communities. As the climate warmed at the end of each ice
age, great successional changes took place. The tundra vegetation and
bare glacial till deposits underwent succession to mixed deciduous forest.
The greenhouse effect resulting in increase in temperature is likely to bring
profound Allogenic changes in the next century. Geological and climatic
catastrophes such as volcanic eruptions, earthquakes, avalanches,
meteors, floods, fires, and high wind also bring allogenic changes.
Mechanisms[edit]
In 1916, Frederic Clements published a descriptive theory of succession
and advanced it as a general ecological concept.[9] His theory of
succession had a powerful influence on ecological thought. Clements'
concept is usually termed classical ecological theory. According to
Clements, succession is a process involving several phases:[9][page needed]

1. Nudation: Succession begins with the development of a bare site,

called Nudation (disturbance).[9]
2. Migration: refers to arrival of propagules.[9]
3. Ecesis: involves establishment and initial growth of vegetation.[9]
4. Competition: as vegetation becomes well established, grows, and
spreads, various species begin to compete for space, light and
nutrients.[9]
5. Reaction: during this phase autogenic changes such as the buildup of
humus affect the habitat, and one plant community replaces
another.[9]
6. Stabilization: a supposedly stable climax community forms.[9]
Seral communities[edit]
Main article: Seral community

Pond succession or sere A: emergent plant life B: sediment C: Emergent plants grow inwards,
sediment accretes D: emergent and terrestrial plants E: sediment fills pond, terrestrial plants
take over F: trees grow

A hydrosere community
A seral community is an intermediate stage found in an ecosystem
advancing towards its climax community. In many cases more than one
seral stage evolves until climax conditions are attained.[20] A prisere is a
collection of seres making up the development of an area from non-
vegetated surfaces to a climax community. Depending on the substratum
and climate, different seres are found.

Changes in animal life[edit]

Succession theory was developed primarily by botanists. The study of
succession applied to whole ecosystems initiated in the writings of Ramon
Margalef, while Eugene Odum’s publication of The Strategy of Ecosystem
Development is considered its formal starting point.[21]
Animal life also exhibit changes with changing communities. In lichen stage
the fauna is sparse. It comprises few mites, ants and spiders living in the
cracks and crevices. The fauna undergoes a qualitative increase during
herb grass stage. The animals found during this stage include nematodes,
insects larvae, ants, spiders, mites, etc. The animal population increases
and diversifies with the development of forest climax community. The fauna
consists of invertebrates like slugs, snails, worms, millipedes, centipedes,
ants, bugs; and vertebrates such as squirrels, foxes, mice, moles, snakes,
various birds, salamanders and frogs.

Microsuccession[edit]
Succession of micro-organisms including fungi and bacteria occurring
within a microhabitat is known as microsuccession or serule. Like in plants,
microbial succession can occur in newly available habitats (primary
succession) such as surfaces of plant leaves, recently exposed rock
surfaces (i.e., glacial till) or animal infant guts,[14] and also on disturbed
communities (secondary succession) like those growing in recently dead
trees or animal droppings. Microbial communities may also change due to
products secreted by the bacteria present. Changes of pH in a habitat
could provide ideal conditions for a new species to inhabit the area. In
some cases the new species may outcompete the present ones for
nutrients leading to the primary species demise. Changes can also occur
by microbial succession with variations in water availability and
temperature. Theories of macroecology have only recently been applied
to microbiology and so much remains to be understood about this growing
field. A recent study of microbial succession evaluated the balances
between stochastic and deterministic processes in the bacterial
colonization of a salt marsh chronosequence. The results of this study
show that, much like in macro succession, early colonization (primary
succession) is mostly influenced by stochasticity while secondary
succession of these bacterial communities was more strongly influenced by
deterministic factors.[22]

Climax concept[edit]
According to classical ecological theory, succession stops when the sere
has arrived at an equilibrium or steady state with the physical and biotic
environment. Barring major disturbances, it will persist indefinitely. This end
point of succession is called climax.
Climax community[edit]
Main article: Climax community
The final or stable community in a sere is the climax community or climatic
vegetation. It is self-perpetuating and in equilibrium with the physical
habitat. There is no net annual accumulation of organic matter in a climax
community. The annual production and use of energy is balanced in such a
community.
Characteristics[edit]

 The vegetation is tolerant of environmental conditions.

 It has a wide diversity of species, a well-drained spatial structure, and
complex food chains.
 The climax ecosystem is balanced. There is equilibrium between gross
primary production and total respiration, between energy used from
sunlight and energy released by decomposition, between uptake of
nutrients from the soil and the return of nutrient by litter fall to the soil.
 Individuals in the climax stage are replaced by others of the same kind.
Thus the species composition maintains equilibrium.
 It is an index of the climate of the area. The life or growth forms indicate
the climatic type.
Types of climax[edit]
Climatic Climax
If there is only a single climax and the development of climax
community is controlled by the climate of the region, it is termed as
climatic climax. For example, development of Maple-beech climax
community over moist soil. Climatic climax is theoretical and develops
where physical conditions of the substrate are not so extreme as to
modify the effects of the prevailing regional climate.
Edaphic Climax
When there are more than one climax communities in the region,
modified by local conditions of the substrate such as soil moisture,
soil nutrients, topography, slope exposure, fire, and animal activity, it
is called edaphic climax. Succession ends in an edaphic climax
where topography, soil, water, fire, or other disturbances are such
that a climatic climax cannot develop.
Catastrophic Climax
Climax vegetation vulnerable to a catastrophic event such as a
wildfire. For example, in California, chaparral vegetation is the final
 vegetation. The wildfire removes the mature vegetation and
decomposers. A rapid development of herbaceous vegetation follows
until the shrub dominance is re-established. This is known as
catastrophic climax.
Disclimax
When a stable community, which is not the climatic or edaphic climax
for the given site, is maintained by man or his domestic animals, it is
designated as Disclimax (disturbance climax) or anthropogenic
subclimax (man-generated). For example, overgrazing by stock may
produce a desert community of bushes and cacti where the local
climate actually would allow grassland to maintain itself.
Subclimax
The prolonged stage in succession just preceding the climatic climax
is subclimax.
Preclimax and Postclimax
In certain areas different climax communities develop under similar
climatic conditions. If the community has life forms lower than those
in the expected climatic climax, it is called preclimax; a community
that has life forms higher than those in the expected climatic climax
is postclimax. Preclimax strips develop in less moist and hotter areas,
whereas Postclimax strands develop in more moist and cooler areas
than that of surrounding climate.
Theories[edit]
There are three schools of interpretations explaining the climax concept:

. Monoclimax or Climatic Climax Theory was advanced by Clements (1916)

and recognizes only one climax whose characteristics are determined
solely by climate (climatic climax). The processes of succession and
modification of environment overcome the effects of differences in
topography, parent material of the soil, and other factors. The whole area
would be covered with uniform plant community. Communities other than
the climax are related to it, and are recognized as subclimax, postclimax
and disclimax.
. Polyclimax Theory was advanced by Tansley (1935). It proposes that the
climax vegetation of a region consists of more than one vegetation
climaxes controlled by soil moisture, soil nutrients, topography, slope
exposure, fire, and animal activity.

. Climax Pattern Theory was proposed by Whittaker (1953). The climax

pattern theory recognizes a variety of climaxes governed by responses of
species populations to biotic and abiotic conditions. According to this theory
the total environment of the ecosystem determines the composition,
species structure, and balance of a climax community. The environment
includes the species responses to moisture, temperature, and nutrients,
their biotic relationships, availability of flora and fauna to colonize the area,
chance dispersal of seeds and animals, soils, climate, and disturbance
such as fire and wind. The nature of climax vegetation will change as the
environment changes. The climax community represents a pattern of
populations that corresponds to and changes with the pattern of
environment. The central and most widespread community is the climatic
climax.
The theory of alternative stable states suggests there is not one end point
but many which transition between each other over ecological time.

Forest succession[edit]

The forests, being an ecological system, are subject to the species

succession process.[23] There are "opportunistic" or "pioneer" species that
produce great quantities of seed that are disseminated by the wind, and
therefore can colonize big empty extensions. They are capable of
germinating and growing in direct sunlight. Once they have produced
a closed canopy, the lack of direct sun radiation at the soil makes it difficult
for their own seedlings to develop. It is then the opportunity for shade-
tolerant species to become established under the protection of the
pioneers. When the pioneers die, the shade-tolerant species replace them.
These species are capable of growing beneath the canopy, and therefore,
in the absence of catastrophes, will stay. For this reason it is then said
the stand has reached its climax. When a catastrophe occurs, the
opportunity for the pioneers opens up again, provided they are present or
within a reasonable range.
An example of pioneer species, in forests of northeastern North America
are Betula papyrifera (White birch) and Prunus serotina (Black cherry), that
are particularly well-adapted to exploit large gaps in forest canopies, but
are intolerant of shade and are eventually replaced by other shade-
tolerant species in the absence of disturbances that create such gaps.
Things in nature are not black and white, and there are intermediate
stages. It is therefore normal that between the two extremes of light and
shade there is a gradient, and there are species that may act as pioneer or
tolerant, depending on the circumstances. It is of paramount importance to
know the tolerance of species in order to practice an effective silviculture.