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Primary Succession

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Primary Succession
Lawrence R Walker, School of Life Sciences, University of Nevada, Las Vegas, Nevada, USA
Roger del Moral, University of Washington, Washington, USA
Primary succession is the assembly of ecosystems on bar-
ren landscapes following severe disturbances that leave
little or no biological legacy (lava flows, landslides and
mine wastes). The assembly process involves colonisation
of newly exposed substrates and subsequent interactions
among the colonising plants, animals and soil microbes.
Successional trajectories of sequential community
replacement then develop, but some may diverge from
nearby trajectories, making succession a challenging
process to predict. Understanding how plant, animal and
microbial communities develop under such extreme con-
ditions is essential for restoration of damaged lands, as
restoration is fundamentally the manipulation of succes-
sional trajectories by humans. Successional studies also
provide insights into loss of biodiversity, climate change
and the influences of invasive species on community
dynamics.
Introduction
Primary succession is a sequential change in species com-
position and other ecosystem characteristics on surfaces
with little or no biological legacy. It can occur following
disturbances such as volcanoes, landslides or mining and
also can occur in aquatic environments on newly exposed
rocks, reefs and shorelines (Walker, in press). Primary
succession concerns humans because it addresses such
fundamental issues as the recovery of plants and animals
and the development of soils following severe disturbances.
Soil erosion is a worldwide phenomenon exacerbated by
human activities that continue to remove essential topsoils
thus threatening the sustainability of agriculture (Pimentel
eLS subject area: Ecology
How to cite:
Walker, Lawrence R; and Moral, Roger del (August 2011) Primary
Succession. In: eLS. John Wiley & Sons, Ltd: Chichester.
DOI: 10.1002/9780470015902.a0003181.pub2
eLS & 2011, John Wiley & Sons, Ltd. www.els.net
Advanced article
Article Contents
. Introduction
. Disturbances that Initiate Primary Succession
. Models of Succession
. Development of Soils
. Factors Influencing Colonisation
. Species Interactions
. Successional Patterns
. Applications
Online posting date: 15th August 2011
et al., 1995) and contributing to global warming by trans-
ferring organic matter to the atmosphere (Conant et al.,
2011). Human activities are also causing other forms of
land degradation. Urbanisation destabilises slopes, creates
more runoff, lowers water tables and covers the landscape
with impervious surfaces. Construction of transportation
corridors has similar effects and further fragments the
landscape. Channelisation of rivers increases the frequency
of their catastrophic flooding. Mining creates extreme
conditions for primary succession by removing topsoil or
forming toxic wastes. Humans also damage freshwater and
marine environments directly through dredging and filling
and indirectly through pollution. Global warming may
intensify hurricanes and coastal flooding and alter climate
patterns. Natural disturbances such as volcanoes, earth-
quakes and dune formation also continue to shape the
landscape and create conditions for primary succession.
Primary succession measures changes that occur over
about 2–1000 years, so it falls between studies of changes in
the fossil record and seasonal fluctuations in plants and
animals (Walker and del Moral, 2003). However, this time
frame indicates a bias towards trees that live several cen-
turies. Succession of short-lived microbes or insects can
occur within hours or weeks. Therefore, change is relative
to the life span of the species of interest. Spatial scales vary,
encompassing the range from test tubes to biomes. Primary
succession also occupies one extreme of a continuum of
disturbance severity and may transition gradually into
secondary succession if soils and some species survive the
disturbance intact. A forest fire may destroy the trees and
understory plants whereas seeds survive, resulting in sec-
ondary succession. But if the fire also destroys all the soil
organisms and burns the organic matter, the situation is
best categorised as primary succession. Similarly, a land-
slide normally removes the topsoil, but if a patch of intact
soil rafts down, settling in the middle of the landslide, a
mosaic of habitats develops (Figure 1). This kind of
patchiness is normal, even with lava, which is perhaps the
most severe disturbance type. Lava flows can leave islands
of intact vegetation or kipukas untouched. It is most useful
to simply describe the levels of soil nutrients and organic
matter, the presence of seeds and vegetative propagules or
1
 Primary Succession
Figure 1 A 6-month-old landslide in the tropical rainforest of the Luquillo
Experimental Forest in Puerto Rico. Photo by Lawrence R Walker.
other physical or biotic conditions rather than worry about
defining the type of succession. Likewise, subsequent dis-
turbances can alter primary succession in ways that are not
severe enough to restart primary succession, so confusion
as to when primary succession actually ends and secondary
succession begins is best avoided by simply explaining the
effects of the relevant disturbances.
Disturbances that Initiate Primary
Succession
Disturbances are relatively discrete events in time and
space that alter the structure of populations, communities
and ecosystems. They can alter the distribution and
amount of biomass or the physical environment (Pickett
and White, 1985; Willig and Walker, 1999). Disturbance,
like primary succession, is a relative term, best defined
within the temporal and spatial scales of interest. It has
characteristics such as origin (from within or outside the
system of interest), frequency, extent (area affected) and
magnitude (intensity of physical force and severity of
damage). The disturbance regime is the combination of all
disturbances within a given area and time and is a function
of the probability of occurrence of each type of
2 eLS & 2011, John Wiley & Sons, Ltd. www.els.net
disturbance. Disturbances often interact, creating a patchy
landscape caused by variations in disturbance severity or
frequency. For example, disturbances such as earthquakes,
volcanoes or road construction destabilise slopes and can
trigger landslides. Landslides can dam rivers and promote
flooding. Volcanoes can melt glaciers and cause flooding
and block rivers to form lakes. The study of primary suc-
cession is best considered at a landscape scale where
establishment is the net result of the disturbance regime on
populations of plants and animals.
Disturbances that create conditions for primary suc-
cession can be grouped by what causes them (e.g. the
classical elements of earth, air, fire and water, (in addition
to those caused by humans). A variety of substrates result
and these vary in stability, texture and fertility. Earth
movements that expose barren substrates include earth-
quakes, volcanoes, landslides, uprooting of trees and
exposure of rock outcrops (Walker, 1999). Fascination
with volcanoes has led to many studies of primary suc-
cession, particularly following eruptions of Krakatau in
1883 and Mount St Helens in 1980 (del Moral and Grishin,
1999). These studies suggest that the trajectories of primary
succession on volcanoes are determined largely by chance
events in a specific context, making outcomes difficult to
predict (del Moral et al., 2009). Landslides and more
gradual forms of soil erosion result from natural disturb-
ances and human activities such as overgrazing. Primary
succession on landslides depends on substrate stability and
organic matter. Air movement can uproot trees and expose
mineral soil for primary succession. Hurricanes disrupt
large areas by direct wind damage, flooding, erosion and
wave damage to coral reefs and shorelines. Dunes also are
wind-derived surfaces where primary succession is limited
by instability, infertility or drought. Fire usually causes
secondary succession unless damage to soils is complete, as
may result from searing volcanic blasts. Water creates
primary surfaces as a liquid and as a solid. Flooding dis-
tributes silts and sands over floodplains where primary
succession is again dependent primarily on substrate sta-
bility and water availability. Glaciers scour the earth’s
surface and when they melt, moraines are subject to pri-
mary succession. Intriguingly, glacial moraines may not
begin as sterile substrates because of active microbial
communities that survived under, in or on top of the ice
(Matthews, 1992).
Models of Succession
Studies of succession have evolved along several con-
trasting themes. Clements (1928) saw succession as a
deterministic process of development with a predictable
trajectory and a stable endpoint or climax. He clearly
linked succession and abiotic disturbances, but thought
that the effects of species on their environment (reaction)
along with other biotic interactions such as competition
were most important once succession began (Walker and
del Moral, 2003). His views were widely influential and

helped establish succession as a central theme in ecology.
However, his analogy of the development of successional
communities with the development of an organism was
sharply criticised. Nevertheless, a holistic view that stems
from Clements is reflected in later efforts to compare suc-
cession to cybernetic systems or derive general principles to
predict temporal change in ecosystem variables (Odum,
1969).
Several contemporaries of Clements (Warming, Cowles
and Gleason) considered succession an indeterminate
process whereby the success of individuals to colonise and
grow at a new site determined the course of succession. This
reductionistic approach arose from an emphasis on the
spatial variability seen in nature and a refusal to generalise
across sites and attribute emergent properties to com-
munities (McIntosh, 1985; Glenn-Lewin et al., 1992). This
approach currently has the most support. However,
Clements’ emphases on the role of disturbance and species
effects on the environment remain important and there is
renewed support for these aspects of succession.
Models of succession are both conceptual and math-
ematical and most apply to both secondary and primary
succession (Glenn-Lewin et al., 1992). Conceptual models
focus on the relative importance of several processes in
determining species change. The successional effects of life
history characteristics such as arrival times, growth rates
and longevities can be modelled for individual plant and
animal species. Interspecific interactions may accelerate
(facilitate) or delay (inhibit) the rate of succession. Some
models attempt to predict succession by using transit-
ion probabilities from prior conditions (e.g. Markov
models), although such models do not address ongoing
environmental fluctuations (Gibson et al., 1997). Models of
forest succession that depend on the properties of indi-
vidual trees mostly have been applied to secondary suc-
cession, but are relevant to later stages of primary
succession. Models have been published that address par-
ticular types of primary succession (Walker and del Moral,
2003) but few attempts have been made to compare pro-
cesses among them. Some progress is being made using
models of alternative stable states that are separated tem-
porally by relatively abrupt transitions (Suding and Hobbs,
2009; Walker and del Moral, 2009). Other recent advances
include the examination of nutrient ratios during suc-
cession (stoichiometric models; Marleau et al., 2011) that
simplify complex issues including spatial heterogeneity of
nutrients and changing patterns of nutrient uptake (Bishop
et al., 2010). Prediction of successional trajectories still
remains most accurate at specific locations, but the
importance of chance events and high spatial variability
confound efforts to generalise about primary succession
(Marteinsdottir et al., 2010).
Development of Soils
The study of primary succession encompasses soil for-
mation and change because most primary seres (a sere is the
eLS & 2011, John Wiley & Sons, Ltd. www.els.net
Primary Succession
sequence of stages in succession) begin without soil. Soil
formation depends on complex interaction of climate,
parent material, topography, organisms and time. Yet
because soil forms slowly relative to the length of human
life spans, few of these variables can be measured directly so
space-for-time substitutions called chronosequences must
be used. However, the assumption in using a chronose-
quence is that the older sites had the same history as the
younger sites. This assumption is often wrong (Johnson
and Miyanishi, 2008), so direct observations over time are
best. Chronosequences are, nevertheless, an important tool
for understanding plant succession at decadal to millennial
time-scales when clear evidence is present that sites of dif-
ferent ages are following the same trajectory. This situation
is most likely when successional trajectories are con-
vergent, have low biodiversity, rapid species turnover and
low frequency and severity of disturbance (Walker et al.,
2010). Trajectories that are divergent, species-rich, highly
disturbed or arrested are less appropriate for chronose-
quence studies.
Soils form most quickly in warm and wet climates where
high primary productivity leads to rapid accumulation of
nutrients and organic matter. Such conditions also favour
rapid decomposition by soil invertebrates, fungi and bac-
teria. Plants influence soil formation through root exud-
ates, root growth, nutrient cycling, water uptake, mineral
weathering, soil aeration and their effects on soil water
content (Bardgett, 2005). Recent advances in measure-
ments of soil microbial function, biomass, respiration and
taxonomy have shown that bacteria usually dominate early
primary succession and fungi dominate later stages
(Ohtonen et al., 1999). Soil microbes may control the
decline in productivity in late stages of primary succession
(retrogression) by their influence on nutrient availability
(Peltzer et al., 2010). Bacteria and fungi, as well as other soil
organisms undergo succession at generally shorter time
scales than plants and are major determinants of the type
and rate of response of soil formation to subsequent
disturbances.
The accumulation of soil nutrients is closely linked to
overall site fertility that determines the quality of leaf litter.
Bacterial fixation of atmospheric nitrogen is especially
important in primary succession where nitrogen levels may
initially be very low. Free-living nitrogen fixers are
important in severe habitats such as polar and temperate
deserts, on recent lava flows, sand dunes or on glacial
moraines. Cryptogamic crusts containing mosses, lichens,
fungi, algae and bacteria can be important stabilisers of
young soils and provide a source of nitrogen (Miles and
Walton, 1993). The largest input of nitrogen is from bac-
teria living in symbiotic association with roots of vascular
plants. These symbiotic relationships often increase the
rate of soil formation (Walker, 1993). Animal parts and
particularly insect feces can provide a significant fraction of
the nitrogen entering the soil during insect outbreaks and
there can be substantial abiotic sources of N as in precipi-
tation. Nitrogen accumulation rates vary from 27 to
163 kg N ha21 year21 and nitrogen levels generally reach a
3
 Primary Succession
plateau after 100–1000 years at levels between 2000 and
5000 kg N ha21 (Walker, 1993). In contrast, soil phos-
phorus is highest early in succession when phosphate is
exposed through erosion or uplift but both available and
total pools decline over time as phosphorus is weathered
and bound in forms unavailable to plants (Crews et al.,
1995).
Soil organic matter accumulates through succession in
conjunction with soil nitrogen, typically reaching values
45% about 60 years after a disturbance that initiates
primary succession (Matthews, 1992; Walker and del
Moral, 2003). The balance between primary productivity
and decomposition regulates the accumulation of organic
matter. Warm and wet climates promote both productivity
and decomposition, decreasing the rate of organic matter
accumulation. Cool but wet climates promote accumu-
lation because of slower decomposition, but when many
primary seres are compared, there is not always a clear
relation to climatic parameters (Wardle et al., 2004).
Factors Influencing Colonisation
Many of the patterns of primary succession can be
explained by understanding the life histories of the species
involved. Much of the uncertainty in predicting succes-
sional trajectories occurs during colonisation. Predict-
ability of colonisation is greatest for small, infrequent
disturbances that are not too severe. Events after colon-
isation are usually more predictable because once estab-
lished, early colonists tend to grow (subject to species
interactions and recurrent disturbances) until they are
either crowded out or die. Any biological legacy due to
survival of seeds, plants and safe sites for establishment
increases predictability (Walker and del Moral, 2003; del
Moral and Eckert, 2005).
Colonisation of newly exposed substrates depends on
landscape factors such as the size and shape of the dis-
turbance as well as biotic factors. These include the com-
position of vegetation surrounding the areas of the new
disturbance and the dispersal abilities of those species
(Glenn-Lewin et al., 1992). Barriers to dispersal include
mere distance (Lanta and Lepš, 2009) and lack of effective
dispersal mechanisms, particularly if widely dispersed
pioneer species are absent from the local flora. If disturb-
ances have been frequent or locally common, pioneer
species may still be well represented and colonisation of the
newly disturbed site will be relatively prompt. The timing of
a gap relative to the production of seeds or vegetative
propagules will also determine which species arrive. Wind,
water or animals can passively carry out dispersal. Birds
and bats are responsible for colonisation to remote areas,
whereas ants and mammals are more important locally.
Humans now provide a nearly global mixing of plants,
especially through introductions for horticulture, agri-
culture and forestry but also through inadvertent intro-
ductions in ballast or cargo on ships and airplanes.
4 eLS & 2011, John Wiley & Sons, Ltd. www.els.net
Plants that survive the disturbance in situ have a jump on
colonisation. Even volcanic disturbances may leave some
plants intact, only lightly buried or with viable seeds.
Vegetative reproduction can result in rapid expansion by
diffusion onto landslides, floodplains, dunes or other
unstable surfaces by plants rooted in stable substrates at
the periphery. Few species are adapted to long distance
dispersal, although large disturbance gaps must initially be
colonised by such infrequent events. Once a surface is
colonised, future generations of colonists are likely to be
controlled by local seed production or vegetative expan-
sion and the disturbance may be colonised in successively
expanding nuclei of plants. This combination of initial long
distance dispersal followed by local diffusion appears to be
the most common method of colonisation in primary suc-
cession (Walker and del Moral, 2003).
Germination and growth must occur for successful col-
onisation. The extreme conditions in primary succession
prevent most dispersed plants from establishing. However,
dispersal of any organic matter including wind-dispersed
arthropods and seeds promotes soil development and
habitat amelioration. Safe microsites for germination
increase over time from both biotic inputs and physical
weathering that provides heterogeneity to the primary
surface. Safe sites can also decline in number as they
become occupied by successful colonists. A safe site must
provide relief from drought or flooding, extreme tem-
peratures and wind and provide nutrients for growth.
Substrate stability is also critical but colonisation of pri-
mary seres may still take decades. When subsequent dis-
turbances occur, succession may be delayed indefinitely.
Species Interactions
The interactions of species alter trajectories and rates of
succession, and primary succession provides a good
opportunity to study the effects of interactions without the
encumbrances of species from previous communities
(Walker and del Moral, 2003). Species interactions are
probably unimportant only in habitats with persistent and
severe disturbances. Species interact along a continuum
from mutualism that provides reciprocal benefits to com-
petition that provides reciprocal harm. One-way effects are
also common, as when species A harms species B but B has
no effect on A. Indirect effects can also influence succession,
as when species A harms C that had harmed B, so B benefits
from the presence of A. Plant species are the most common
focus of succession studies, but plant interactions with soil
organisms determine soil formation and plant interactions
with animals such as herbivores also impact both plant and
animal succession.
Connell and Slatyer (1977) described three models of
succession that addressed the importance of species inter-
actions to species replacements. Their tolerance model
emphasised the lack of interactions early in succession,
followed by survival of species with the most tolerance of
declining fertility and light. Their facilitation model

followed Clements’ idea that the first colonists improved
the habitat for successive colonists but not for self-repro-
duction. There is much support for this from primary
succession, where the initial environment is too harsh for
many species to colonise. However, the facilitation is not
considered obligatory because many species do well in
certain favourable microsites (Callaway and Walker,
1997). Their inhibition model suggested that early colonists
inhibited later arrivals by monopolising available
resources. Connell and Slatyer (1977) found most support
for this model in secondary succession, but further studies
have confirmed that competitive inhibition is also common
in primary succession (Matthews, 1992). Even large vas-
cular plants with nitrogen-fixing bacterial symbionts that
are considered potential facilitators because of their con-
tribution of nitrogen and high quality litter can form dense
thickets that inhibit successive colonists (Walker, 1993).
During the life span of a single plant there can be both
facilitative and inhibitory interactions with other plants
(Walker et al., 2003). The net effect on succession comes
from the sum of all possible interactions in any successional
stage and has not yet been determined effectively for any
single case.
The relative importance of facilitation and inhibition
varies along stress and productivity gradients. In severe
environments, large plants may benefit smaller plants or
later colonists by providing protection from wind, desic-
cation, strong light or herbivory. In less severe environ-
ments, the net effect of the larger nurse plant may actually
be inhibitory by competing with the smaller plant for
nutrients, light or water (Callaway and Walker, 1997). The
relationship of competition and facilitation to productivity
is less clear, with evidence that competition and facilitation
increase, decrease or remain constant with increases in
productivity in different cases (Walker and del Moral,
2003). Comparative studies of primary seres along prod-
uctivity gradients will help determine whether any patterns
exist for this relationship. However, because competition
and facilitation can have different effects on the various life
history stages of a plant, the focus must remain on the net
effect of the myriad interactions.
Successional Patterns
The consequence of colonisation and species interactions
on developing soils is a particular trajectory of species
change that occurs at a given rate. Trajectories can con-
verge towards or diverge from the species composition of
the mature, undisturbed vegetation or from patterns
occurring on similar disturbances. Convergence occurs
when initial differences due to the stochastic nature of
colonisation and environmental heterogeneity are over-
ruled by biologically based processes or there is a signifi-
cant biological legacy (del Moral, 2007). Where
competitive inhibition is strong or where species pools are
small due to a stressful environment, convergence is likely.
eLS & 2011, John Wiley & Sons, Ltd. www.els.net
Primary Succession
Divergence occurs when the random events of colon-
isation lead to differences in vegetation that in turn persist
by competitive inhibition. Heterogeneity of local soil or
environmental conditions can also produce divergence,
particularly if it develops after the initial disturbance (del
Moral et al., 2010). Divergence may also arise if there is a
large species pool available for colonisation of later stages
of succession. Spatial patchiness or a mosaic of land use
patterns and disturbances can lead to divergence because
both physical and biological processes are rarely syn-
chronous across the landscape. Local convergence may
arise from dominance by a competitive species, but diver-
gence can still exist at larger scales (Lepš and Rejmánek,
1991). Other types of trajectories include parallel,
deflected, arrested, retrogressive or cyclical trajectories, but
convergence and divergence are most likely (Matthews,
1992; Walker and del Moral, 2003).
The rate of succession determines when the landscape
regains its pre-disturbance functions (Anderson, 2007).
Restorationists, farmers and wildlife managers are all
concerned with ecosystem function for different reasons.
Rates can be measured as the accumulation of vegetative
cover, biomass or species richness, as the rate of change of
species (turnover) or by the time required to reach a certain
stage. Both abiotic and biotic factors control the rate of
successional change. When physical resources such as light,
water and nutrients are adequate, biotic interactions
determine rates of change. The initial conditions of the site,
subsequent disturbances and the vagaries of colonisation
also affect successional development.
Patterns of community and ecosystem development
during primary succession are as varied as those of sec-
ondary succession. To move beyond simple generalisations
that soils develop, nutrients and organic matter accumu-
late, biomass accumulates and structural complexity
increases, we need more studies that use comparable
methods to evaluate difference among and between types of
primary succession. Predictability of trajectories and rates
remains elusive, but the expansion of long-term permanent
plot studies will allow direct measurements of the par-
ameters of successional change and lessen reliance on
indirect chronosequence methods.
Applications
Primary succession provides a model for guiding restor-
ation of damaged lands and can provide insights into loss
of biodiversity, climate change and the role of invasive
species (Prach and Walker, 2011). Natural processes of
recovery can take longer than is desirable for restoration
programs and conditions may have become so degraded
that thresholds of irreversibility have been crossed so that
succession will not occur without intervention. Therefore,
integration of both scientific and management concerns is
needed, the former for an understanding of basic mech-
anisms and generalities, the latter for practical knowledge
5
 Primary Succession
about methods, local concerns and societal issues (Walker
et al., 2007).
Because successional trajectories remain substantially
unpredictable, restoration goals will often go unmet if they
focus on particular sequences of species replacements.
Instead, more general expectations of increased soil
organic matter, plant biomass or plant cover can be sub-
stituted. In areas of low biodiversity or harsh environments
species able to colonise may be limited and succession more
predictable. However, attempts to repeat natural suc-
cession are unlikely to succeed. One problem is that in
highly modified environments such as urban areas, seed
sources are unlikely to arrive at a disturbance without
assistance. This issue can be resolved with artificial intro-
ductions of seeds and seedlings. Secondly, and more chal-
lenging, shortcuts to the final stage or even linear recreation
of various stages with dominant species is also unlikely to
replicate natural succession. Too many unknowns such as
the role of soil biota, understory plants, animal pollinators
and herbivores make simple replacement of a few plant
species a poor substitute for natural processes. With the
urgency of most restoration attempts demanding faster-
than-normal ecosystem recovery, recreation of only some
aspects of succession is expected. Rehabilitation, or any site
improvement, is achievable, but restoration, or the com-
plete establishment of an original ecosystem, is unrealistic.
Interactions between successional theory and practical
applications in restoration ecology can provide a positive
feedback loop that improves land management practices
(del Moral et al., 2007). Stabilisation of substrates and
removal of surface compaction increase the probability of
initiating succession. Fertilisers and mulches can amelior-
ate harsh conditions and improve plant survival. Initial
plantings of grasses or aggressive herbs may stabilise sub-
strates in the short term but inhibit establishment of later
successional species. Managers often learn these lessons
from trial and error, but scientists can help determine the
degree of manipulation needed at a given site and the use-
fulness of lessons learned at other sites. Natural conditions
result in many endpoints for primary succession and
ecologists can provide alternative scenarios to managers.
In some cases, novel communities may produce the best
results.
There are many ways that successional studies can help
to explain loss of biodiversity. Monitoring of biodiversity
and experimental manipulations during succession suggest
that peaks in biodiversity often occur in early but not initial
stages of succession, and that populations are vulnerable at
different life history and successional stages to extirpation
from disturbances (Wardle et al., 1999; Isbell et al., 2009).
Successional studies can also determine current trends in
biodiversity loss and help predict future patterns (Vellak
et al., 2009; Prach and Walker, 2011).
Successional studies can help clarify climate change
implications through long-term monitoring of rates of
species change, reanalysis of historical data and experi-
mental manipulations such as additions of carbon dioxide
(Smith et al., 2000; Karlsen and Elvebakk, 2003; Parmesan,
6 eLS & 2011, John Wiley & Sons, Ltd. www.els.net
2006; Prach et al., 2010). Finally, successional studies can
elucidate effects of invasive species through analyses of
colonisation (invasion) patterns across disturbance and
fertility gradients (Meiners et al., 2007), examination of
competitive interactions among species (Yurkonis et al.,
2005), and clarification of the role of invasive species in
creating novel ecosystems (Hobbs et al., 2006).
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Dynamics and Restoration. Washington, DC: Island Press.
Johnson EA and Miyanishi K (2007) Plant Disturbance Ecology:
The Process and the Response. Amsterdam: Academic Press.
7
 Primary Succession

Luken JO (1990) Directing Ecological Succession. London:
Chapman and Hall.
del Moral R and Walker LR (2007) Environmental Disasters,
Natural Recovery and Human Responses. Cambridge: Cambridge
University Press.
Tilman D (1988) Plant Strategies and the Dynamics and Structure
of Plant Communities. Princeton, NJ, USA: Princeton Uni-
versity Press.

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