Received: 20 December 2018 | Accepted: 14 January 2019

DOI: 10.1111/1365-2745.13132

EDITORIAL

Ecological succession in a changing world

Cynthia C. Chang1 | Benjamin L. Turner2

1
Division of Biology, University of
Washington, Bothell, Washington Abstract
2
Smithsonian Tropical Research Institute, 1. Ecological succession – how biological communities re‐assemble and change over
Balboa, Republic of Panama
time following natural or anthropogenic disturbance – has been studied since the
Correspondence birth of ecology, and the resulting theoretical framework underpins many aspects
Cynthia C. Chang
of the discipline. Recently, the mechanistic basis of classic succession theory has
Email: cynchang@uw.edu
been advanced by studies of plant and microbial interactions, functional traits,
Handling Editor: Richard Bardgett
and retrogressive stages of ecosystem development. This special issue brings to‐
gether a series of papers that highlight these contemporary novel approaches and
how our understanding of ecological succession has advanced.
2. Four key themes emerge from the issue: (a) generalizations about succession, (b)
the influence of dispersal and habitat size on successional trajectories, (c) changes
in plant functional traits during succession, and (d) belowground community inter‐
actions during long term during ecosystem development.
3. Synthesis. The articles in the special issue highlight novel perspectives on succes‐
sion theory, revealing the importance of historical contingency, disturbance sever‐
ity, dispersal limitation, functional traits, and belowground community processes
in determining patterns of ecosystem development. Together, they reinforce the
importance of ecological succession in understanding the response of plant and
microbial communities to disturbance in a changing world.

KEYWORDS
community assembly, dispersal limitation, disturbance, disturbance severity, global change,
ecosystem development, functional traits

1 | I NTRO D U C TI O N and has direct relevance to studies of landscape ecology, ecosystem

Ecological succession – the study of how biological communities re‐as‐
semble following natural or anthropogenic disturbance – has been a
foundation in ecology, and the theoretical framework underpins many
aspects of the discipline (Egerton, 2015; Meiners, Cadotte, Fridley,
Pickett, & Walker, 2014; Prach & Walker, 2011; Walker & Wardle,
2014). Although succession is sometimes perceived as an old‐fash‐
ioned topic, recent studies and reviews demonstrate that succession
continues to play a central role in modern ecological theory and ap‐
plication. For example, our understanding of succession is embedded
in theories of modern community assembly and species coexistence
(Chang & HilleRisLambers, 2016; HilleRisLambers, Adler, Harpole,
Levine, & Mayfield, 2012; Pulsford, Lindenmayer, & Driscoll, 2014),
development, restoration ecology, and global change ecology (Meiners
et al., 2014; Prach & Walker, 2011; Walker, Walker, & Hobbs, 2007;
Walker & Wardle, 2014). In particular, successional studies provide
insight into the community assembly mechanisms, including dispersal
limitation (Makoto & Wilson, 2016; Tilman, 1993), species pool effects
(Li et al., 2016), priority effects (Fukami, 2015), abiotic environmen‐
tal filtering (Lebrija‐Trejos, Perez‐Garcia, Meave, Bongers, & Poorter,
2010; Lohbeck et al., 2014), stochastic processes (Marteinsdóttir,
Svavarsdóttir, & Thórhallsdóttir, 2018; Norden et al., 2015), biotic in‐
teractions (e.g. competition, facilitation, herbivory; Connell & Slatyer,
1977; Huston & Smith, 1987; Tilman, 1993), and feedbacks (e.g. be‐
tween plant and microbial communities; Bauer, Mack, & Bever, 2015).
Ecological succession clearly serves as a foundation for modern ecology.

Journal of Ecology. 2019;107:503–509. wileyonlinelibrary.com/journal/jec

© 2019 The Authors. Journal of Ecology | 503
© 2019 British Ecological Society
504 | Journal of Ecology
Recent advances in our understanding of ecological succession
are particularly relevant in the current era of rapid global change.
For example, land‐use change and habitat fragmentation directly af‐
fect dispersal, species pool sizes, and priority effects (e.g. Damschen
et al., 2008; De Meester, Vanoverbeke, Kilsdonk, & Urban, 2016;
Fukami, 2015). Changes abiotic conditions due to nutrient deposi‐
tion or climate change affect environmental filtering and the strength
and direction of plant–soil feedbacks, with potential consequences
for community development (e.g. Blois, Zarnetske, Fitzpatrick, &
Finnegan, 2013; Ettinger & HilleRisLambers, 2013; Härdtle, Niemeyer,
Niemeyer, Assmann, & Fottner, 2006). Furthermore, because natural
and anthropogenic disturbances (e.g. fire, volcanic eruption, recla‐
mation sites, and old‐field/forest recovery from land‐use) influence
community assembly mechanisms in different ways, they provide
valuable natural experiments to disentangle the mechanisms of com‐
munity change. Studies of ecological succession can therefore help
understand how ecosystems respond to global change.
Recently, the application of modern techniques and data analy‐
ses have advanced our understanding of ecological succession. For
example, developments in molecular analysis have allowed the study
of succession in belowground microbial communities over both short
and long timescales (Jangid, Whitman, Condron, Turner, & Williams,
2013; e.g. Bauer et al., 2015; Cutler, Chaput, & van der Gast, 2014),
while functional trait measurements (e.g. Diaz & Cabido, 2001;
Garnier et al., 2004) and null model analyses (e.g. Dini‐Andreote,
Stegen, van Elsas, & Salles, 2015; Purschke et al., 2013) have provided
novel perspectives on community assembly. Updated conceptual
frameworks have also advanced our understanding of succession.
For example, primary succession is now understood to extend be‐
yond “climax” communities, as ecosystems continue to develop into a
retrogressive phase linked to long‐term depletion of soil phosphorus
(Peltzer et al., 2010; Wardle, Walker, & Bardgett, 2004). These novel
approaches have revealed mechanisms that drive succession, and
addressed key questions such as when stochastic versus determinis‐
tic processes dictate succession trajectories (Li et al., 2016; Norden
et al., 2015) and how abiotic factors versus biotic interactions influ‐
ence these community and ecosystem outcomes (Fridley & Wright,
2012; HilleRisLambers et al., 2012). Only by examining succession
across a variety of ecosystems can we begin to understand whether
there are universal rules governing the process of succession.
Originating from an Organized Symposium in August 2017 at
the 102nd Annual Meeting of the Ecological Society of America in
Portland, Oregon, this special issue brings together a series of articles
that highlight how contemporary studies are bringing novel perspec‐
tives to our understanding of ecological succession. The articles span a
range of disturbance regimes and timescales during both primary and
secondary succession. They examine successional patterns in plant and
belowground microbial communities in a diverse range of ecosystems,
including temperate and tropical grasslands and forests, sub‐alpine vol‐
canic landscapes, and human‐disturbed reclamation sites. Four main
themes emerge (Figure 1): (a) generalizations about succession, (b) the
role of dispersal and habitat size in driving successional trajectories, (c)
functional trait dynamics during succession, and (d) the influence of
EDITORIAL
belowground community interactions on long‐term ecosystem devel‐
opment. Together, the articles extend our conceptual framework for
understanding succession by providing novel insights into the mecha‐
nisms driving community assembly in ecosystems worldwide.
1.1 | Generalizations about succession
Despite succession being a foundation of ecological theory, few studies
have sought broad generalizations across a range of successional sites.
Such comparative studies are necessary because they provide insight
into the relative importance of community assembly mechanisms (e.g.
environmental filtering, biotic interactions, priority effects, and dispersal
limitation) at different stages of succession (Chang & HilleRisLambers,
2016), in different successional types (e.g. primary versus secondary
succession), and across broad spatial and temporal scales (Walker, 2011;
Walker & Wardle, 2014). Broad comparative studies also provide the
opportunity to understand successional trajectories, including whether
communities recover to a previous state or diverge to a new state (Prach
et al., 2016), which provides a basis for understanding and informing
restoration management (Suding & Hobbs, 2009; Walker & del Moral,
2009). This special issue contains four papers that specifically address
the theme of generalizations in succession.
Understanding how patterns of succession differ across a dis‐
turbance severity gradient (e.g. primary versus secondary succes‐
sional sites) provides context for how disturbances and their severity
influence community trajectories and recovery rates. By explicitly
comparing primary and secondary succession across a broad range
of ecosystems in the published literature, Prach and Walker (2019)
report that primary successional sites more often increased in spe‐
cies richness, had more divergent trajectories, and suffered less im‐
pact of alien species compared to secondary succession sites. Their
meta‐analysis also reveals that recovery rates differ among ecosys‐
tem types, for instance between cold and warm biomes, suggesting
that global climate changes will influence community recovery rates
and restoration management practices in the future.
There have been few opportunities to compare the effect of
disturbance across a severity gradient within a single ecosystem.
Taking advantage of the variety of disturbance impacts following
the Mount St. Helens (Washington, USA) volcanic eruption in 1980,
Chang et al. (2019) synthesized three 36‐year datasets to examine
succession patterns across a disturbance gradient ranging from pri‐
mary to secondary succession. They found that disturbance severity
played a greater role in succession than small‐scale influences such
as site history and local environment, yet, surprisingly, communi‐
ties were most variable in the intermediately disturbed, secondary
successional sites. Sites suffering high and low‐severity disturbance
recovered more slowly, but in response to different mechanisms
(seed/abiotic conditions versus light limitation, respectively). These
findings indicate that rates of community change are driven not
only by disturbance severity, but also by the various mechanisms
that mediate community dynamics. Together, the studies of Prach
and Walker (2019) and Chang et al. (2019) suggest that it is possi‐
ble to make broad generalizations about disturbance severity and
EDITORIAL
F I G U R E 1 Themes in ecological succession, and the methods used to advance theory and knowledge in basic and applied ecology in a
rapidly changing world [Colour figure can be viewed at wileyonlinelibrary.com]
succession, although confirmation of driving mechanisms requires
further research.
Understanding the patterns and rates of succession is critical
to determining whether the recovery of plant communities follows
a particular successional trajectory after disturbance, yet there are
few opportunities to experimentally compare recovery rates in dis‐
turbed and non‐disturbed communities at a large scale. In this issue,
Fischer, Antos, Biswas, and Zobel (2019) examined the influence of
disturbance severity following the eruption of Mount St. Helens by
harnessing a remarkable long‐term experiment in which tephra de‐
posits were removed from understory forest plots in the immediate
aftermath of the eruption. They found that recovery was site‐spe‐
cific, because initial conditions and disturbance intensity interacted
to determine community change. Similarly, Clark, Knops, and Tilman
(2019) found that divergent successional trajectories in the herba‐
ceous community after 88 years of old‐field succession were due
to contingent (non‐random) factors such as site conditions, com‐
petition, and demographic trade‐offs rather than stochastic (ran‐
dom) factors. To identify which of these major drivers influenced
successional dynamics, the authors compared meta‐community
model simulations to long‐term old‐field succession data at Cedar
Creek, Minnesota, USA. They discovered that compensatory trade‐
offs between colonization and mortality rates drive the divergent
patterns of community change found over the course of succession.
Together these studies highlight the importance of deterministic
mechanisms such as disturbance intensity, site conditions, competi‐
tion, and demographic trade‐offs on successional trajectories.
1.2 | Influence of dispersal limitation and habitat
size on succession
A number of factors influence which species are able to colonize and
persist as community assembly occurs over the course of succes‐
sion (HilleRisLambers et al., 2012). For example, successional trajec‐
tories reflect regional species pool size, distance from seed source,
and functional traits related to dispersal (Jones & del Moral, 2009;
Makoto & Wilson, 2016; Tilman, 1993), while habitat size affects
Journal of Ecology | 505
species pool size and environmental conditions at local scale (e.g.
space availability, increased biotic interactions). An understanding
of these community assembly mechanisms during succession has
importance in understanding meta‐community dynamics, landscape
ecology, invasion ecology, and restoration ecology. This special issue
has three papers that explore the role of dispersal limitation and
habitat size on successional processes.
Many past studies have focused on the role of dispersal limita‐
tion on early successional stages (Walker & del Moral, 2003), while
studies on late successional stages often focus on ecosystem de‐
velopment (e.g. Wardle et al., 2004). In a meta‐analysis of primary
succession sites across many systems, Makoto and Wilson (2019)
examined the role of dispersal limitation on both the early and late
stages of succession. They found that dispersal limitation influ‐
ences rates of succession in both the early and late stages, even
after centuries, and overall appears to limit key processes linked to
ecosystem development such as vegetation cover and soil carbon
accumulation. In a study of secondary succession in neotropical
forests, van Breugel et al. (2019) found that both dispersal limita‐
tion and soil nutrient status caused variation in species composition
at local and landscape scales, although the influence of soil nutri‐
ents fades during later successional stages, presumably reflecting
increasing light limitation as forests mature. Their findings indicate
that spatial heterogeneity in the landscape can promote resilience
of ecological communities in human‐modified ecosystems.
Habitat size can also influence species pool size and site condi‐
tions, and both mechanisms directly impact community assembly.
By studying a gradient of island size caused by habitat fragmen‐
tation created by a dammed river in China, Liu et al. (2019) found
that island size influenced functional trait composition, diversity,
and successional direction, and that rates of succession were faster
on large compared to small islands. Overall, habitat size influenced
successional patterns more than isolation effects in a study system
where seed dispersal is facilitated predominantly by birds, and veg‐
etation regeneration occurs in the context of secondary succession.
Together, these studies have implications for generaliza‐
tions about successional theory, and for the applied contexts of
506 | Journal of Ecology
community response to global climate change and habitat fragmen‐
tation. For example, climate change will affect dispersal processes
in direct (e.g. dispersal phenology, traits) and indirect ways (e.g. en‐
vironmental factors, biotic interactions), further influencing succes‐
sional trajectories (Makoto & Wilson, 2019). Similarly, understanding
human land use and habitat fragmentation will require insight into
factors that drive secondary successional processes and community
recovery following land abandonment. The importance of habitat
size and site conditions highlighted in this issue therefore provides
critical insight into restoration and land management plans.
1.3 | Functional trait dynamics over the
course of succession
Plant traits that influence fitness or performance provide a mech‐
anism to understand patterns of plant distribution and productiv‐
ity in the environment (Diaz & Cabido, 2001; Garnier et al., 2004;
Reich, 2014; Wright et al., 2004, 2010). In turn, this has significance
for understanding successional patterns by providing insight into
species turnover through environmental filtering during succes‐
sion (Kraft et al., 2014; Lasky, Uriarte, Boukili, & Chazdon, 2014).
There has been considerable interest in patterns of plant traits dur‐
ing succession (Douma, de Haan, Aerts, Witte, & van Bodegom,
2012), with a number of recent studies in grasslands (e.g. Kahmen &
Poschlod, 2004; Kelemen et al., 2017; Purschke et al., 2013), forests
(e.g. Chai et al., 2015; Derroire, Powers, Hulshof, Varela, & Healey,
2018; Lasky et al., 2014; Lohbeck et al., 2014), and volcanic islands
(Karadimou et al., 2018). Two new studies extend this for grasslands
and tropical forests.
Traits related to nutrient dynamics are linked to plant metabo‐
lism and can vary markedly among species, thus representing a key
aspect of functional diversity. To examine factors influencing the
relative success of invasive plant species in temperate grasslands,
Duffin, Li, and Meiners (2019) quantified foliar nutrients as func‐
tional traits in native and exotic taxa in New Jersey, USA, and linked
this to information on long‐term community dynamics. Successional
trajectories of abundance‐weighted foliar nutrients differed mark‐
edly between native and exotic species, suggesting that the success
of exotic invasive species is influenced by selection of plant traits
during succession. In tropical forests of Singapore, Lai, Chong, Yee,
Tan, and van Breugel (in revision)1 examined the extent to which
plant functional traits influence recruitment during secondary suc‐
cession in treefall gaps. Although traits such as specific leaf area ap‐
peared unimportant, they found evidence that wood density, seed
mass, and adult stature all influence the development of the sapling
community. These results suggest that traits related to the leaf eco‐
nomics spectrum have little influence on recruitment following dis‐
turbance, with successional trajectories instead explained by
multiple life history tradeoffs. Together, they add to a growing body
of evidence showing how functional trait change can help under‐
stand community dynamics during ecosystem development.
1
Accepted articles published in Journal of Ecology but appearing after the Special Issue.
EDITORIAL
1.4 | Influence of belowground community
interactions on succession
There is an increasing interest in successional patterns in the soil mi‐
crobial community, both for understanding how heterotrophic and
symbiotic organisms change over time, and how these changes link
to corresponding changes in plant communities. A number of studies
have examined this topic, across short and long timescales, on a variety
of parent materials and under a variety of climates (e.g. Jangid et al.,
2013; Tarlera, Jangid, Ivester, Whitman, & Williams, 2008; Uroz, Tech,
Sawaya, Frey‐Klett, & Leveau, 2014). While past studies have explored
belowground microbial communities, they have typically focused on
a single taxonomic group (most often the bacteria), but advances in
metagenomics now allow for broader integration across taxonomic
groups, including assessment of symbiotic organisms and linkages
between plant and microbial communities and plant–soil feedbacks.
Three papers in the special issue specifically address this theme.
Turner et al. (2019) examined patterns of α‐diversity for archaea,
bacteria, and fungi over 2 million years of ecosystem development
along the Jurien Bay chronosequence of coastal dunes in Western
Australia. The chronosequence is of particular interest because it
represents an extremely strong fertility gradient in a global plant
diversity hotspot, allowing the assessment of coordination in the re‐
sponse of plants and soil microbial communities to long‐term pedo‐
genic change. The authors found contrasting patterns of diversity in
biological communities, with the greatest α‐diversity of prokaryotes
on young or intermediate‐aged soils, but eukaryote diversity increas‐
ing continuously throughout the chronosequence. Although plant
and microbial communities became increasingly decoupled through
time, the changes for both groups were linked to soil acidification
during pedogenesis, revealing a coordinated driver of above and
below ground diversity during long‐term ecosystem development.
The community composition of mycorrhizal fungi shifts during
succession (e.g. Bauer et al., 2015; Gao et al., 2018; Johnson,
Zak, Tilman, & Pfleger, 1991), which might influence successional
patterns through positive plant–soil feedbacks (Bever, Platt, &
Morton, 2012). In this issue, Koziol and Bever (2019) examined the
influence of arbuscular mycorrhizal fungi on ecological succession
in prairie grasslands. By growing plants in mesocosms inoculated
with fungi from different successional stages, they found that early
successional plants consistently exhibited negative frequency–de‐
pendent growth, while late successional plants demonstrated pos‐
itive frequency–dependent growth in the presence of beneficial
fungal partners. These findings were supported by results from
field inoculation assays, in which beneficial fungi facilitated late
successional plant establishment. The results indicate that posi‐
tive plant–mycorrhizal feedbacks accelerate plant community suc‐
cessional trajectories once late successional plants establish.
Mycorrhizal symbioses vary predictably as soil nutrient availabil‐
ity changes from nitrogen to phosphorus limitation, involving a shift
from plant communities dominated by arbuscular mycorrhizal plants
on relatively young soils, to an increasing abundance of ectomycor‐
rhizal and ericoid mycorrhizal plants on older soils (Lambers, Raven,
EDITORIAL
Shaver, & Smith, 2008; Zemunik, Turner, Lambers, & Laliberté,
2015). Although most species form associations with only one type
of fungus, some species can associate with either arbuscular or ec‐
tomycorrhizal fungi, and often form associations with both types
simultaneously (e.g. Dickie, Koide, & Fayish, 2001; McGuire et al.,
2008). Furthermore, some of these plant species associate with ni‐
trogen‐fixing bacteria, forming a triple symbiosis. In this issue, Teste
and Laliberté (2019) examined the symbiotic associations of two
dual‐infected species, one of which also associates with nitrogen‐
fixing bacteria, across a long‐term chronosequence of coastal dunes
in Western Australia. They found plasticity in symbiotic associations
during long‐term ecosystem development, although the “favoured”
symbiont predominated on both young and old soils despite marked
variation in soil fertility. In addition, by simultaneously assessing the
influence of biotic effects (through inoculation) and abiotic condi‐
tions (along the chronosequence), they demonstrate the importance
of soil properties in driving patterns of plant and root responses
during long‐term ecological succession.
Collectively, these three studies highlight the remarkable insight
that belowground communities can bring to our understanding of
ecological succession. Given our relatively limited understanding
of microbial community dynamics during succession, and the rapid
advances being made in metagenomic analysis, it seems likely that
this topic will continue to yield important novel insight into the
mechanistic underpinnings of ecological succession in the coming
decade.
2 | FU T U R E C H A LLE N G E S
This special feature identifies several opportunities for future re‐
search that will improve our understanding of the drivers and pat‐
terns of succession. First, the papers in this special issue highlight
the need for large temporal and spatial scales to best account for
inherent variation across these scales, which provides the basis for
understanding the extent to which successional dynamics can be
generalized. This highlights the value of long‐term datasets, chron‐
osequences, and large‐scale studies to understanding successional
processes, and we encourage efforts to maintain and sample long‐
term study sites along successional gradients. Second, papers in this
issue demonstrate that combining experimental manipulation with
long‐term monitoring can provide unique insight into the specific
mechanisms that drive successional patterns. Matching large‐scale
and/or long‐term community change patterns with mechanisms that
explain these patterns are critical to allowing general succession
theory to be used in more applied restoration and management set‐
tings. Finally, the articles in this special issue reveal novel ways in
which studies of succession can inform ecosystem and community
responses to climate change, habitat fragmentation, and restoration
ecology. Such connections between ecological theory and applied
ecology are fundamental to understand the challenges facing eco‐
systems in our rapidly changing world. As such, we hope that that this
special issue not only highlights the continued value and relevance
Journal of Ecology | 507
of successional research, but also encourages future successional
research to be better integrated into relevant global change issues.
AC K N OW L E D G E M E N T S
We thank the Ecological Society of America for hosting the sym‐
posium on Ecological Succession, Dr. John Bishop for his con‐
tribution to the organization of the symposium, Dr. Lawrence
Walker for critical comments on the manuscript, and Prof. Richard
Bardgett and James Ross for their support in organizing and edit‐
ing the special issue.
DATA AC C E S S I B I L I T Y
No data were used for this editorial piece.
ORCID
Cynthia C. Chang https://orcid.org/0000-0002-3281-4864
Benjamin L. Turner https://orcid.org/0000-0002-6585-0722
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How to cite this article: Chang CC, Turner BL. Ecological
succession in a changing world. J Ecol. 2019;107:503–509.
https://doi.org/10.1111/1365-2745.13132