Succession
JM Pandolﬁ, University of Queensland, Brisbane, QLD, Australia
r 2008 Elsevier B.V. All rights reserved.
Introduction
This article explores the notion of ecological succession, one of ecology's oldest and most controversial concepts. Simply stated,
succession is the orderly changes in communities of a speciﬁc place over a period of time. More formally, it is the orderly
progression of directional community development that results from modiﬁcation of the physical environment by the community
and culminates in an ecosystem in which maximum biomass and interactions among component organisms are maintained.
Many workers relate succession to disturbance (ﬁres, hurricanes, drought, etc.), such that succession refers to changes in a
community after a perturbation opens up ecospace. Most of the earlier work on succession (dating back to 1899!) described the
sequential invasion of species invading a particular site; later studies deal with a whole range of community characteristics from
both empirical and theoretical standpoints (Table 1). Most of the work on succession has been dominated by forest communities,
but successional studies also exist for a number of other ecosystems including benthic and pelagic marine ecosystems.
Different Kinds of Succession
Two kinds of succession have been proposed. `Autogenic' succession refers to sequential changes in the community that are
brought about by the inﬂuence and activities of the organisms themselves upon the environment or habitat. `Allogenic' succession
occurs to communities through the inﬂuence of factors external to the organisms themselves. The utility of allogenic succession
may be limited because it may be too broadly applicable to any temporally repetitive change. Most of the debates surrounding
succession, and studies which provide detailed examples are concerned with autogenic succession.
`Primary' succession refers to the colonization of sites where a severe disturbance has left no trace of the preexisting community,
or where entirely new ecospace has been created. These might include a terrestrial or submarine lava ﬂow, glacial recession, or the
movement of submarine or subaerial sand dunes. Harsh environments usually characterize the earliest phases of primary succession,
and communities tend to develop more slowly than with more minor disturbances to the ecosystem. `Secondary'
succession refers to colonization of sites which did have a previous community established, but a perturbation has removed a
portion of it or all of it over a limited area. Here again, much of the debate surrounding succession and most of its examples come
from secondary succession. However, most unassailable instances of succession are examples of primary succession.
Expected Community Trends of Succession
The expected community trends of succession are explicit and detailed (Table 1). They rely on an early-successional community
that ultimately leads to the `climax' community, or the most stable state for the successional sequence. This climax community is
maintained until an extrinsic disturbance resets the successional clock. The attributes are grouped according to community
energetics, community structure, life-history characteristics, cycling of nutrients, selection pressure, and overall homeostasis. These
trends follow from the notion that the climax community is a steady state maintained by internal feedback control mechanisms.
But they break down if it is not, and there is little evidence so far that they are.
In forests there are a variety of aspects of production ecology which vary over the successional sequence including forest ﬂoor
depth, chemistry and content of decaying wood, nutrient availability, leaf area, carbon allocation, species composition and their
nutritional adaptations, the relative roles of trees and minor vegetation, the relative importance of geochemical, biogeochemical
and internal cycles, rate of soil organic matter accumulation, and total ecosystem organic matter. Early-successional forest species
generally are shade intolerant, require a mineral seedbed, may be fast-growing, can generally utilize nutrients that occur in newly
formed gaps, may be nitrogen-ﬁxing, are tolerant of microhabitats in newly disturbed settings, and are superior competitors in
such settings. Late-successional forest species generally are shade tolerant where they are able to grow slowly, may require a more
organic seedbed, and are generally poorly adapted to newly disturbed settings where they are poor competitors. Nutrients are
generally gained from litter decomposition and mycorrhizae.
Models of Succession
Three alternative models have been proposed for determining the sequence of species that might occur after a perturbation. The
ﬁrst is the facilitation model in which only certain species (`early successional') are able to arrive immediately following a
Encyclopedia of Ecology, 2nd edition, Volume 3 doi:10.1016/B978-0-444-63768-0.00547-3616
disturbance. The most important factor governing succession in the facilitation model is the degree to which early-successional
species alter the environment for later-successional species. Later species can only colonize after these early-successional species
have sufﬁciently modiﬁed the environmental conditions. For example, in forest communities the amount of net primary production
of the plant community will inﬂuence its ability to alter the site. This model implies a high degree of organization in
ecological communities. However, it may still operate in communities where such organization is not apparent, for example, in
heterotrophic successions or primary successions.
The second is the tolerance model in which the sequence of species to inhabit an ecosystem after a disturbance is determined
exclusively by their life-history characteristics. The species that occur are simply those that are most efﬁcient in exploiting resources
in a given time or place. Thus, the success of later species is unrelated to the species composition of the earlier community. Two
circumstances have been forwarded where this model of succession might be most prevalent. The ﬁrst is with certain groups of
animals, such as large vertebrates or insects that display a high degree of social integration, that have evolved a high degree of
independence from both biotic and physical environments. The second is in situations where populations are limited more by
resources than natural enemies or environmental stress. In the tolerance model, life-history characteristics such as the relative
longevity, growth form, reproductive mode, and size of the early versus later species becomes paramount. A further elaboration of
this model is given below in the section titled `The role of competition and life history'.
The third is the inhibition model in which early-successional species inhibit the success and growth of later-successional
species. Those species that happen to occupy the site ﬁrst maintain their membership in the community until their populations are
damaged or die, only then enabling later species to invade. As such, the species composition gradually shifts to species with greater
longevities. No special mechanism need be invoked for the inhibition model of succession.
Most models of succession have been criticized. Some authors treat them as purely descriptive and, like the entire successional
process, a particular manifestation of the outcome of interacting individual life histories. In fact, one of the original authors of
these three models has recently acknowledged that directional species replacement is only one change that might occur after a
disturbance, and is often not the most likely change.
Table 1 Predicted changes that occur in major structural and functional characteristics of a developing ecosystem during ecological succession
Ecosystem attributes Developmental stages Mature stages
Community energetics
1. Gross production/community respiration (P/R ratio) Greater or less than 1 Approaches 1
2. Gross production/standing crop biomass (P/B ratio) High Low
3. Biomass supported/unit energy ﬂow (B/E ratio) Low High
4. Net community production (yield) High Low
5. Food chains Linear, predominantly grazing Weblike, predominantly detritus
Community structure
6. Total organic matter Small Large
7. Inorganic nutrients Extrabiotic Intrabiotic
8. Species diversity – variety component Low High
9. Species diversity – equitability component Low High
10. Biochemical diversity Low High
11. Stratiﬁcation and spatial heterogeneity (pattern diversity) Poorly organized Well-organized
Life history
12. Niche specialization Broad Narrow
13. Size of organism Small Large
14. Life cycles Short, simple Long, complex
Nutrient cycling
15. Mineral cycles Open Closed
16. Nutrient exchange rate, between organisms and environment Rapid Slow
17. Role of detritus in nutrient regeneration Unimportant Important
Selection pressure
18. Growth form For rapid growth (`g-selection') For feedback control (`K-selection')
19. Production Quantity Quality
Overall homeostasis
20. Internal symbiosis Undeveloped Developed
21. Nutrient conservation Poor Good
22. Stability (resistance to external perturbations) Poor Good
23. Entropy High Low
24. Information Low High
Analysis of these parameters provides empirical tests of the expected development of ecosystems during succession.
Reprinted with permission from Odum EP (1969) The strategy of ecosystem development. Science 164: 262–270. Copyright 1969 AAAS.
General Ecology: Succession 617
Some authors have taken these three models and related them to the search for ecological assembly rules during succession and
compared them with a fourth alternative, the random colonization model, where chance survival of different species initiates the
successional sequence; subsequent associations occur as a result of subsequent random colonization by new species. The search for
assembly rules in ecology has a rich and controversial history that parallels that in succession. In fact, successional changes in
community composition might be viewed as a special case of an overall search for deterministic processes in the maintenance and
diversity of ecological communities.
Factors Inﬂuencing Succession
There are a number of factors that can inﬂuence the way in which community succession proceeds, if at all. Probably the most
important of all are differences in life-history traits such as dispersal, ability to establish in open sites, shade tolerance, etc. The
tradeoff between the availability of seeds and propagules of later-successional species versus the degree to which the earlysuccessional
community is resistant to invasion will inﬂuence the degree to which the former can increase at the successional site.
Invasion success itself is dependent upon a large number of factors including, but not limited to, changes in microclimate or
microhabitats, the composition of the seedbed, diseases, predators, and competition. Superimposed upon all of these are factors
external to the organisms themselves, such as precipitation, temperature, salinity, and disturbance regime which can serve to reset
the successional sequence.
The Role of Competition and Life History
Patterns from succession are numerous and varied, and have been mainly reported from the population and ecosystem level. However,
a generalized theory of succession has been proposed for plants based on individual organisms. It is based on a model with three
features: (1) the demographic history (birth, growth, death) of each individual is tracked through time; (2) every individual is assigned a
suite of species-speciﬁc life-history traits (e.g., size, maximum age, and shade tolerance); and (3) competition and resource availability
(light) is modeled explicitly through varying each individual's growth and mortality based on its degree of shade tolerance and light
availability. The model successfully produces a signiﬁcant portion of the variability in successional patterns using the mechanism of
competition. Again, the importance of life-history traits among interacting plants is manifested.
Two-species simulations from the model were classiﬁed into ﬁve groupings based on the temporal patterns of species
divergence (Fig. 1). The ﬁrst was classical successional replacement where one species was able to dominate early in succession
and the other species later, and is due to the correlation among life-history parameters of the two species. Divergence
occurs when competitive ability starts out the same, but then one species outcompetes the other whereas convergence is the
opposite: species are unequal in competitive ability early in succession but this equals out later in succession. Total suppression
occurs when one species has the competitive advantage throughout the entire successional development. Finally,
pseudocyclic replacement occurs when a temporary period of dominance or co-dominance by the initial cohort of species 2
results during the period between the senescence of the initial cohort of species 1 and the maximum lifespan of species 2.
Thus early succession is characterized by the establishment of both species, but the initial cohort of species 2 persists longer
than that of species 1. Species 2 is later replaced by the competitively superior species 1. The outcomes of the simulations
depend more upon the equality of the life-history characteristics among species as opposed to the absolute values of the
characteristics themselves; thus outcomes are governed by the qualitative relationship between the species with respect to
their competitive abilities through time. Results of the simulations were similar to the two-species models when more than
two species were considered and when multiple resources were considered. To summarize this important model to the
concept of succession, the summary of the authors is quoted:
An approach based on competition among individual plants is presented as an explanation for species replacements during plant succession. Inverse
correlations among life-history and physiological traits that confer competitive ability under different environmental conditions are shown to be
sufﬁcient to produce successional replacements but not sufﬁcient for understanding the complex variety of successional patterns unless they are
applied at the individual rather than at the population level or higher. With models based on competition among individual plants, various
combinations of life-history and physiological traits can produce the great variety of population dynamics found in natural successions. The classic
successional pattern of species replacement results from a particular structure of correlations among life-history sand physiological characteristics.
Atypical patterns of succession result when this correlation structure is altered. Both primary and secondary succession are modeled as nonequilibrium
processes capable of interacting with disturbances to produce steady-state communities whose properties depend on abiotic conditions, such as
temperature and resource levels, and on the type and frequency of disturbances. (Huston and Smith, 1987, p. 193)
Before leaving the subject of competition, it must be noted that there may well be a role for positive interactions among
species comprising a community. There is increasing evidence that direct positive interaction, or facilitations, among species can
play a vital role in ecological communities. While relatively unexplored in successional studies, the role of facilitations might
provide an important context for testing the importance of successional patterns and processes in the maintenance of
communities.
618 General Ecology: Succession
Is Succession Real?
Many models of succession are based on the idea that species that occur early in the successional sequence facilitate their own
replacement by altering the environment to be more favorable to later species. Of course, this is anathema to natural selection – no
species would `intentionally' create more favorable circumstances for any other species than itself (the `evolutionary paradox' of
succession). For this reason the whole idea of succession has been questioned, and some authors consider it to be no more than
the result of the passage of time, with no real mechanistic basis.
Many of the factors that might inﬂuence succession have been called into question, such that many authors believe that it is not
a real phenomenon, or if it is, is unimportant in the maintenance of species diversity. Of critical importance is the degree to which
communities act in changing the environment. Some reviews of the idea of succession conclude that most purported examples of
succession do not ﬁt the predicted trends: changes in structural and functional properties are not consistently associated with
changes in species composition; successional stages are not consistently unidirectional; and effects of earlier species might just as
easily delay or prevent as facilitate later-successional replacement.
Workers critical of successional theory have also put forward alternatives. One of these relates purported successional changes
to consequences of differential growth, survival, and colonizing ability of species adapted to growth at different points on
environmental gradients. Differences in taxonomic composition among successional stages are dependent upon interspeciﬁc
competition while structural and functional changes in the community results in correlations in plants among size, longevity, and
growth rate. In fact, successional changes may be exclusively governed by the presence of slower-growing later-successional species
during the early phases of succession.
Much of the debate stems from the classic divergence in opinion about the inﬂuence of factors that result in the way in which
species are maintained within ecological communities. On the one hand there may only be a loose association of species who
1
2
1
Biomass
2
(a)
(e)
1
2
(b)
1
2
Time
(c)
1
2
(d)
Fig. 1 General patterns of species interactions in a two-species succession. Competitive dominance occurs as a result of differing life-history
traits. (a) Classical successional replacement that results from an inverse relationship between the two species in the attributes that confer early
and late competitive advantage; (b) divergence that results when competitive abilities are equal early on, followed by a competitive advantage by
one of the species; (c) convergence, the inverse of (b), that results when one species has a competitive advantage that equals out during the
succession; (d) total suppression that results from one species having a competitive advantage over the other throughout succession; and (e)
pseudocyclic replacement that results from alternating periods of dominance or co-dominance between the two species. From Huston MH and
Smith T (1987) Plant succession: Life history and competition. American Naturalist 130: 168–198.
General Ecology: Succession 619
happen to inhabit the same habitat at the same time. In this model, species do not `feel' each other or interact much, or if they do,
the effects of these interactions are minor in comparison with their shared environmental tolerances. In the extreme view,
communities are just haphazard associations of species and any spatial or temporal continuity in the composition of a community
is either minimal, or determined solely by environmental parameters.
A completely different view holds that biological communities are tightly integrated units composed of species that not only
respond together to environmental changes, but also maintain linkages among each other that result in continuity of occurrence
over broad spatial and temporal scales. Some caricatures of this model have even referred to the community as a `super-organism'
with emergent properties that extend beyond individual species. While this view is probably overstated, there is a large amount of
evidence that species interactions within communities result in spatial and temporal continuity of species assemblages within
habitats. It is these interactions and other biological properties that have led many authors to believe that succession is a very
plausible model for understanding changes in species composition over time.
Some Examples of Succession
Regardless of the continuing debates on whether there is any mechanistic basis for explaining directional patterns in species
occurrences, excellent examples of the latter occur in nature. One of the best studies of primary succession was a test of the
mechanisms involved in succession following deglaciation at Glacier Bay, Alaska. The main conclusion from this work was that no
single factor or mechanism can totally account for primary succession. Plant life-history traits, such as seed size, growth rate, age at
ﬁrst reproduction, maximum height, and longevity played a fundamental role, but competition among species, and facilitation by
addition of soil N and organic matter were also important. Although the relative importance of these factors was inﬂuenced by
resource availability and environmental stress, they varied predictably through the successional sequence. The authors concluded
that (1) life-history traits and availability of propagules determine the pattern of succession, (2) the mechanism for changes in
species abundance patterns that accompanies succession is competition, and (3) the rate of change and nature of the climax
community depends upon initial site conditions and the role of facilitation.
Perhaps one of the most famous and long-standing successional stories comes from the Krakatau Islands of Indonesia. The
volcanic eruptions between May and August 1883 resulted in the almost complete `sterilization' of the islands. A classic primary
successional sequence has been documented for the main volcanic island of Rakata which continues uninterrupted, whereas the
other islands display characteristics of secondary succession (Fig. 2). On Rakata, a number of different successional pathways are
being expressed (Fig. 2a). A similar diversity of succession characterizes the other islands, but eruptive activity of Anak Krakatau – a
new volcano that was established in the center of the group between 1927 and 1930 – disrupted the vegetation of some of the
other islands such as Serung and R. Kecil. There, varying disturbance events have reset the successional sequence (Fig. 2b).
Differences in vegetation and in the curves of species colonization among the Krakatau Islands have been attributed mainly to
intermittent disturbances of varying intensities from Anak Krakatau. The importance of succession in viewing the history of
terrestrial ecosystems of the Krakatau Islands is summed up by Whittaker et al.
Neither the pattern of change in the ﬂora, nor that for some groups of fauna, e.g., the birds and butterﬂiesy can be understood without reference to
vegetation succession and the key period in the 1920s when the savanna vegetation of the interiors gave way to forest. y a successional model of
island recolonization is required, involving evaluation of habitat and dispersal mechanisms. (Whittaker et al., 1989, p. 103)
Succession in the Fossil Record
One of the great limitations of assessing the frequency or validity of the processes of succession is that the complete successional
sequence might take longer than the period of time over which most ecosystems are studied within scientiﬁc research programs.
Since succession is by its very nature a process that occurs over time, it is not surprising that paleontologists have investigated the
degree to which it can be understood from evidence in the fossil record. In contrast to modern ecologists working on mainly
terrestrial forest communities, paleoecologists have focussed more on marine systems, studying succession in fossil coral reefs,
marine hardgrounds, and some sandy substrates. A critical debate for paleoecologists is the degree to which species replacements
in communities are ecological versus evolutionary phenomena.
Palaeontologists have used ancient reefs as model ecosystems in the study of succession. Four major phases of succession in
eight ancient reefs ranging in age from the Early Ordovician (488 Ma) to the Late Cretaceous (65.5 Ma) have been established:
stabilization, colonization, diversiﬁcation, and domination. The stabilization stage involves the initial colonization of the seaﬂoor
that results in the establishment of a ﬁrm substrate. The colonization stage is characterized by encrusters and frame-builders, those
organisms capable of colonizing a hard substrate and that begin to build three-dimensional structures above the sediment–water
interface. Some authors consider the ﬁrst two of these stages to be the same, equivalent to the pioneering stage of ecologists
studying succession in living forest communities. The third stage in the succession of fossil reefs is referred to as the diversiﬁcation
stage, where maximum diversity of the reef community is developed. Diversity then decreases as a `domination' stage sets in
whereby a single functional entity characterizes the reef community. As the communities ascend through the ﬁrst three stages of
succession, species diversity, degree of stratiﬁcation and pattern diversity, niche specialization, and the complexity of food chains
all increase. These successional sequences typically occur through signiﬁcant intervals of time and are the result of the gradual
620 General Ecology: Succession
alteration of the submarine substratum by individual species, and the elaboration of energy-ﬂow pathways as the community
proceeds through time. Shorter-term successional episodes have also been observed where occasional environmental perturbations
destroy the preexisting community. Here, subsequent community development involves a rapid biologically induced sequence of
communities that culminate in the climax community that was established prior to the perturbation. Just as in the longer-term
successional sequences, increases in major functional and structural attributes of the community accompany these shorter-term
successional sequences.
There are, of course, a number of potential problems associated with the identiﬁcation of succession in the fossil record. In the
model above, an increase in diversity may come about just as easily from an increase in the zonation of the reef as to intrinsic
biological changes. In the domination phase, the accompanying decrease in diversity may reﬂect a community that caps a reef after
it dies, as opposed to a decrease during the life of the reef. However, in the fossil record of the Quaternary period (the past
1.8 Myr) these difﬁculties can often be overcome due to the precise age dating and sea-level history of the reefs.
Pes-caprae
beach
community
(a)
(b) Pes-caprae
beach
community
Pes-caprae and
Casuarina
Terminalia
Barringtonia
forest
Ficus
pubinervis
forest
Neonauclea
forest
Neonauclea
forest
Neonauclea
forest
Neonauclea
forest
Mossy forest
Ficus and
Neonauclea etc.
Schefflera (and Cyrtandra)
scrub with low trees 1983
1951
1932
1919
1906
1897
1886
1883
1983
1908
Sterilization
Inland
Sterilization
Ferns and Cyanophyceae
Ferns and
Cyanophyceae
Ferns and
Cyanophyceae
Ferns and
Cyanophyceae
Pioneer grasses
Pioneer
Pes-caprae
Coastal
Coastal
Pioneer grasses
(Saccharum)
Mature Saccharum
with scattered trees
Cyrtandra
scrub
Cyrtandra?
etc.
Cyrtandra
scrub
Schefflera and Cyrtandra
scrub
Cyrtandra, Saccharum
and ferns
Terminalia
Terminalia
Terminalia
forest
Lower region
(to 550 m)
Upper region
(above 550 m)
Summit
Casuarina
wood
Saccharum
Mature Saccharum
with scattered trees (1919)
1897
1883
Macaranga-Ficus
forest (1920s)
Timonius
Timonius
forest
Dysoxyium
forest
Casuarina
Casuarina
Beachaccretion
Pes-caprae
Macaranga-Ficus
Fig. 2 Principal successional pathways on (a) Rakata and (b) Sertung and R. Kecil plant communities. Rakata has not experienced any volcanic
disturbance for over 100 years, whereas forest development on Sertung and R. Kecil has been interrupted by volcanic activity several times since
1930 by Anak Krakatoa. Eruptive activity changed late-successional forest communities to early-successional or `pioneer' communities. From
Whittaker RJ, Bush MB, and Richards K (1989) Plant recolonization and vegetation succession on the Krakatau Islands, Indonesia. Ecological
Monographs 59: 59–123.
General Ecology: Succession 621
In Kenya, `obligatory succession' in the Pleistocene consisted of an early assemblage of sediment-tolerant corals (dominated by
massive or doming corals) that was replaced by predominantly branching and platy/encrusting corals. This kind of succession was
mainly conﬁned to the earliest portions of reef development on a bare substrate. Patterns through longer successional phases were
varied, but under certain conditions massive or domed-shape corals might replace the branching assemblages. However, clearly
deﬁned zones were rare – most temporal changes in reef species associations were random or unstructured.
Applications to Restoration Ecology
The concepts of succession and community assembly have recently been taken up by the ﬁeld of `restoration ecology'. The nature
of community assembly and the degree to which communities can be expected to return on a path generally directed to their
original states (though return to their exact previous state is probably not possible nor desirable) is of primary interest to
restoration ecologists interested in mitigating and reversing the effects of human-induced habitat degradation. Because of this
central concern and the fundamental questions which each theory provides, it is an important exercise to understand the
similarities and differences between community succession and community assembly. Both concepts recognize the importance of
historical events in shaping community composition, which develops over time toward a ﬁnal state, and both recognize competition
as a major driving force.
Single or multiple stable states are predicted for both concepts, but the way in which species are maintained in such states is
very different. We have already discussed the idea of a climax community – the ﬁnal stable state of a successional sequence.
However, where succession occurs, multiple stable states can be maintained across an ecosystem by disturbances that keep the
community in a variety of successional states, so the `climax' is never achieved at any one site or time. Where these disturbances are
generated by the characteristics of the community itself (herbivory or ﬁre) the community may be kept at an `arrested' successional
state. Arrested succession might also arise where early-successional species maintain their advantage over later-successional species
for extremely long periods of time. `Cyclic' succession occurs when there is no one stable successional state and all of the stages
yield to others within the successional cycle. Stochastic differences in the arrival and colonization of late-successional species might
also result in variability among ecosystems in their community composition.
Community assembly refers to the process by which species colonize, interact with other species and establish a community.
This assembly may or may not be affected by `assembly rules' which predict community composition on the basis of a few key
attributes such as size of the species pool, the abiotic environment, or interspeciﬁc interactions. Community assembly may also
result in either a single community type or multiple stable states. Where there is a strong match between environment and
community, then community composition should converge towards a single state where similar environmental conditions hold,
regardless of the historical order in which the species invade. But multiple stable equilibria have also been observed with varying
historical sequence of species invasions, so that different timing of introduction of species colonization leads to different species
composition.
Regardless of whether species are maintained through a successional sequence, or the product of their sequence of invasion into
an ecosystem, an understanding of the processes which bring about various community states is fundamental to those wishing to
move the system in one direction or another. Even where such processes are particularly opaque, an understanding of the presence
of particular states may help in providing goals for management and restoration. Some predictions about the likelihood of the
existence of a single or multiple stable states have been put forward. While these need further testing, a single stable community
state has been predicted to arise in ecosystems with small species pools, high levels of dispersal, low productivity, and high rates of
disturbance; whereas multiple stable states should be more apparent in ecosystems with large species pools, low levels of dispersal,
high productivity, and low rates of disturbance. Thus, an integrated approach to understanding the processes underlying the
historical trajectory of ecosystems, coupled with an in-depth knowledge of life-history variability, and a few key attributes of the
ecosystem can go far in our understanding of how communities are likely to respond in the future to environmental
changes today.
Further Reading
Chapin, F.S., Walker, L.R., Fastie, C.L., Sharman, L.C., 1994. Mechanisms of primary succession following deglaciation at Glacier Bay, Alaska. Ecological Monographs 64,
149–175.
Clements, F.R., 1916. Plant Succession: An Analysis of the Development of Vegetation. Washington, DC: Carnegie Institute.
Connell, J.H., Slayter, R.O., 1977. Mechanisms of succession in natural communities and their role in community stability and organization. American Naturalist 111,
1119–1144.
Cowles, H.C., 1899. The ecological relations of the vegetation on the sand dunes of Lake Michigan. Botanical Gazette 27, 95–117. 167–202; 281–308; 361–391.
Crame, J.A., 1980. Succession and diversity in the Pleistocene coral reefs of the Kenya coast. Palaeontology 23, 1–37.
Drury, W.H., Nisbet, I.C.T., 1973. Succession. Journal of the Arnold Arboretum 54, 331–368.
Egler, F.E., 1954. Vegetation science concepts. I. Initial ﬂoristic composition, a factor in old-ﬁeld development. Vegetatio 4, 412–417.
Gleason, H.A., 1926. The individualistic concept of the plant association. Bulletin of the Torrey Botanical Club 53, 7–26.
Colonization, Succession and Stability. In: Gray, A.J., Crawley, M.J., Edwards, P.J. (Eds.), The 26th British Ecological Society Symposium held jointly with the Linnean Society
of London. Oxford: Blackwell.
Huston, M.H., Smith, T., 1987. Plant succession: Life history and competition. American Naturalist 130, 168–198.
622 General Ecology: Succession
Lawton, J.H., 1987. Are there assembly rules for successional communities? In: Gray, A.J., Crawley, M.J., Edwards, P.J. (Eds.), Colonization, Succession and Stability: The 26th
British Ecological Society Symposium Held Jointly with the Linnean Society of London. Oxford: Blackwell, pp. 225–244.
McCook, L.J., 1994. Understanding ecological community succession: Causal models and theories, a review. Vegetatio 110, 115–147.
Odum, E.P., 1969. The strategy of ecosystem development. Science 164, 262–270.
Walker, K.R., Alberstadt, L.P., 1975. Ecological succession as an aspect of structure in fossil communities. Paleobiology 1, 238–257.
Whittaker, R.J., Bush, M.B., Richards, K., 1989. Plant recolonization and vegetation succession on the Krakatau Islands, Indonesia. Ecological Monographs 59, 59–123.
Young, T.P., Chase, J.M., Huddleston, R.T., 2001. Community succession and assembly. Ecological Restoration 19, 5–18.
General Ecology: Succession 623