Journal of Vegetation Science 25 (2014) 1113–1116

SPECIAL FEATURE: VEGETATION PATTERNS AND THEIR UNDERLYING PROCESSES

Vegetation patterns and their underlying processes:
where are we now?
Aveliina Helm, Rein Kalamees & Martin Zobel
SPECIAL FEATURE EDITORS

Helm, A. (corresponding author,
aveliina.helm@ut.ee),
Kalamees, R. (rein.kalamees@ut.ee) &
Zobel, M. (martin.zobel@ut.ee): Institute of
Ecology and Earth Sciences, University of
Tartu, Lai 40, 51005 Tartu, Estonia
Abstract
This Special Feature of the Journal of Vegetation Science contains ten contributions
from the 56th Symposium of the International Association for Vegetation Sci-
ence (IAVS), which was held in Tartu, Estonia, 26–30 Jun 2013, and focused on
processes underlying vegetation patterns. These contributions give an overview
of recent advances in vegetation science, with topics ranging from the introduc-
tion of new methods and ideas to comprehensive analysis of vegetation patterns
on a global scale. The focus of these Special Feature contributions indicates that
vegetation science is becoming more global, more complex and more interdisci-
plinary. It is likely that progress in methods and accumulation of data have led
vegetation science into a ‘new era of discovery’.

Overview of 2013 IAVS Symposium
This Special Feature of the Journal of Vegetation Science origi-
nates from the 56th Symposium of the International Asso-
ciation for Vegetation Science (IAVS) held in Tartu,
Estonia, 26–30 Jun 2013. The main theme of the sympo-
sium – Vegetation patterns and their underlying processes
– allowed the incorporation of topics that have attracted
the curiosity of vegetation scientists for the past decades.
The aim of the symposium was not to focus solely on tradi-
tional descriptive and experimental approaches, but rather to
explore new possibilities offered by contemporary molecular
methods, improved computing power and accumulating data
sets that increase our knowledge of global vegetation patterns
and background information. Sessions covered aspects of veg-
etation from global studies to small-scale analysis of below-
ground biota, as well as from historical vegetation patterns to
future novel ecosystems. Altogether, 450 participants from 41
countries gave 374 presentations in 20 oral sessions, four spe-
cial sessions and two poster sessions. Eight insightful plenary
lectures were given by invited keynote speakers, and two
momentous talks were given by award-winners: David Til-
man received the Alexander von Humboldt Medal and deliv-
ered a talk ‘Biodiversity: From evolutionary origins to
ecosystem functioning’, and long-time IAVS member Bastow
Wilson received an IAVS Honorary Membership Award and
gave a lecture entitled ‘Four theories in vegetation science’,
which created lively discussion.
The IAVS 56th Symposium in Tartu was a great success,
and feedback from the participants was overwhelmingly
€
positive (Opik et al. 2014). The organizers thank each and
every delegate for interesting science and camaraderie. Even
the Estonian weather, which is famous for its unpredictabil-
ity, played along, resulting in field excursions with sunshine
and only occasional showers. An alternative and equally
likely option would have been 10 °C and constant rain.
Contributions to this Special Feature
The symposium keynote speakers and organizers of special
sessions were asked to give overviews of their topics,
resulting in ten mostly essay-type contributions to this
Special Feature. These contributions address innovative
theory, methodology and application of vegetation ecology
over a range of topics and scales. For example, Moles et al.
(2014) and Fraser et al. (2014) present global approaches
on plant traits and diversity, whereas Wilson (2014) tackle
issues of below-ground patterns at the local community
level. Similarly broad is the temporal scale of the contribu-
tions – Reitalu et al. (2014) encourage plant ecologists to
look more boldly into the data and methods of Quaternary
palaeoecologists, while Eriksson (2014) provides a
thought-provoking analysis of how anthropogenic pres-
sure in habitats can induce eco-evolutionary dynamics: a
process that can shape the future of our ecosystems.
Until recently, the below-ground part of plants has
received relatively little attention in vegetation science,
due mostly to methodological constraints (P€ artel et al.
2012). Current research, however, indicates that the
below-ground component – i.e. root systems, mycorrhiza,

Journal of Vegetation Science

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Editorial: Special Feature
rhizosphere bacterial communities – can play a significant
role in the development of vegetation patterns and
diversity (van der Putten et al. 2013). Additionally, distri-
bution, richness and ecological relationships of plants
below-ground can differ from those above-ground
(Hiiesalu et al. 2012), which challenges a number of eco-
logical theories and hypotheses that have developed
mostly on the basis of the above-ground component of
vegetation. Recent advances in technology and DNA
sequencing have opened up new sources of information
for vegetation scientists. The paper by keynote speaker
Scott Wilson (2014) in this Special Feature gives an over-
view of how knowledge on the below-ground component
can provide a more complete picture of patterns and func-
tions of vegetation. Minirhizotron technology allows us to
‘take a peek’ at spatial and temporal distribution of organ-
isms underground, whereas DNA sequencing of mixed
species root samples sheds light on below-ground vegeta-
tion composition. Contributions by Mari Moora (2014) –
organizer of the special session on soil biota – and keynote
speaker Martin Zobel (Zobel & Opik € 2014) tackle the role
of mycorrhiza in plant communities. Moora (2014) argues
that possible effects of mycorrhiza in plant communities
are often overlooked by vegetation scientists. To overcome
this shortcoming, she provides an overview of mycorrhiza-
related plant traits that should be considered in vegetation
studies, as well as suggests a novel index to describe the
mycorrhizal trait composition of plant communities – com-
munity mycorrhization. Zobel & Opik € (2014) analyse the
still unanswered question, whether plant communities are
driven by the composition of their mycorrhizal partners
(driver hypothesis) or vice versa, whether mycorrhizal com-
munities are dependent on changes in plant communities
(passenger hypothesis). They show that it might be either,
depending on the successional state of habitats, but also
argue that under stable conditions within-region co-varia-
tion of plant and mycorrhizal communities might depend
mostly on abiotic conditions, and suggest a novel habitat
hypothesis to test this idea.
Drivers of plant community assembly and diversity are
also topics of the Special Feature contributions of Eriksson
(2014), Cerdeira et al. (2014), P€artel (2014), and Fraser
et al. (2014). Keynote speaker Ove Eriksson provides fresh
insight on eco-evolutionary dynamics in plant communi-
ties (2014). Rapid evolution, triggered by novel influences
such as new enemies, new habitats or changing environ-
mental conditions, is noted to occur more often than previ-
ously thought (Buswell et al. 2011). Eriksson (2014)
provides additional insight on a possible feedback mecha-
nism: rapid evolution of species can influence selection via
alteration of ecological processes within communities.
These eco-evolutionary dynamics can be a result of
human-mediated niche construction, and can have
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A. Helm et al.
significant influence on formation of species pools, vegeta-
tion patterns and community assembly.
Issues of niche are also the topic of the contribution
from Cerdeira et al. (2014), who suggest mathematical
solutions to improve geometric procedures in characteriz-
ing niche. The authors revisit Hutchinson’s niche concept,
capturing the mathematical notions implicit in his ideas, to
identify those issues that should be addressed for a closer
link between ecological theory and models. This approach
can significantly improve modelling of ecological pro-
cesses. In the IAVS 2013 Symposium, mathematical niche
construction topics were merged into the special session
‘Niche-based approaches: tackling the link between envi-
ronment and biota’.
Meelis P€ artel (2014) summarizes his plenary talk in a
paper discussing the issue of dark and hidden diversity.
Dark diversity comprises species that are currently absent
in the community, although their ecological suitability and
geographical distribution allows them to be present. Hidden
species are those that are actually present, but are missed
during inventories for various reasons. P€ artel (2014) shows
that knowledge of absent species is vital to understand
community biodiversity and to improve decision-making
in biodiversity conservation. Although intuitive and useful,
this approach is novel to community ecology, brought to
wider attention only in 2011 (P€ artel et al. 2011). No doubt
the ecology of absent species will gain wide use in the
future, as stated by P€artel (2014): ‘species presences have
meaning only if there are absences’.
Two papers in this Special Feature settle for nothing less
than describing the whole world: Fraser et al. (2014) and
Moles et al. (2014) both aim to test classical questions in
ecology with large, global cover data sets. Lauchlan Fraser
et al. (2014) organized the special session ‘Productivity
and Diversity’ and urges ecologists to join the global stan-
dardized sampling protocol HerbDivNet to test the relation-
ship between productivity and species diversity in
grassland communities. The authors of Fraser et al. (2014)
argue that despite recent efforts (e.g. Adler et al. 2011),
this long-debated issue in ecology still lacks sufficient clar-
ity. HerbDivNet aims to put the validity of Grime’s (1973)
hump-backed curve to test through a coordinated
approach with specifically designed methodology.
Keynote speaker Angela Moles et al. (2014) asks
another classical, yet unanswered question: which is more
important for plants, temperature or precipitation? They
use a data set from worldwide joint efforts – the plant trait
data set TRY (Kattge et al. 2011) – and test for the first time
whether global variation in plant trait values depends
more on mean annual temperature or mean annual pre-
cipitation. After a complicated analysis, this simple ques-
tion receives an elegant answer: both are relatively poor in
determining global-scale patterns of plant traits, but
Journal of Vegetation Science

A. Helm et al.
temperature is a significantly better predictor of plant trait
values than precipitation. These kinds of approaches as
presented in Moles et al. (2014) and Fraser et al. (2014),
which put basic ecological hypotheses and theories to com-
prehensive global tests, are likely to become more common
in the future.
In his plenary keynote, Petr Pysek (Pysek & Chytr y and methods, we cannot forget the demand for the under-
2014) addressed plant invasions – an applied topic that also
attracts the attention of theoretical ecologists. Empirical
information on invasion patterns in plant communities
has been accumulating for quite a long time and Pysek &
Chytry emphasize that the time is ripe to analyse the mac-
roecological perspective and community context of plant
invasions. The authors list general topics for which we
have enough empirical information to move towards gen-
eralizations. They also challenge the geographic bias of
invasion studies – most of the empirical information still
comes from Europe, the United States, Australia and New
Zealand.
Although vegetation scientists and Quaternary palaeoe-
cologists deal with similar topics, simply within different
time frames, the representatives of both disciplines rarely
meet. The special session ‘Past vegetation patterns’ held in
IAVS 2013 was considered to be timely and necessary by
both sides. The quality of reconstructions of ancient vege-
tation structure and composition is constantly increasing.
This makes it possible to increasingly apply the same meth-
ods used in conventional plant community ecology and to
address topics that have traditionally not been in the
sphere of interest of palaeoecology – e.g. functional diver-
sity and assembly rules in palaeovegetation (Reitalu et al.
2014).
Vegetation science – where are we heading?
Current advances show that the future will lead us to more
global, more interdisciplinary and also more complex vege-
tation science. Accumulating background data and larger
computing power are taking us from traditional descriptive
and experimental approaches to more holistic and complex
views of ecosystems.
The increasing amount of information on climate, soil
properties and human influence allows the building of bet-
ter models to reveal the underlying factors that shape veg-
etation patterns. Principal hypotheses and theories in
ecology that have not often been properly examined, but
rather rely on small-scale observations or even purely on
dogmatic beliefs, can be tested with ameliorating global
data sets. Standardized and unified methodologies allow
generalizations over different regions and climatic condi-
tions (De Caceres & Wiser 2012; Fraser et al. 2014). Molec-
ular methods and advanced technology will lead to more
thorough exploration of below-ground communities and
Journal of Vegetation Science
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Editorial: Special Feature
help to resolve the role of microorganisms and mycorrhiza
in plant communities, as well as characterize below-
ground vegetation structure and function. All this indicates
that much research is still ahead and the future is full of
opportunities for vegetation science.
However, during the enthusiasm provided by new tools
lying data. It is curious that in an era of vast scientific dis-
coveries and sophisticated methods, we often still lack
basic information on plant distribution patterns even in
relatively well-studied regions such as Europe (Kier et al.
2005). Information from other regions and of other groups
of organisms is even scarcer, including, for instance,
pollinators or fungi that can reveal substantial information
on patterns also shaping vascular plant diversity (Lavorel
et al. 2013). Thus, the need to ‘describe the world’ and
continuously improve our knowledge on distribution and
diversity of species remains. Thorough information on
current vegetation patterns is especially important to track
the effects of global change and human impact on
ecosystems.
We believe that vegetation science is experiencing a
new ‘era of discovery’. There are more possibilities than
ever to acquire a comprehensive understanding of vegeta-
tion patterns and their underlying processes both locally
and globally, and this knowledge is crucial to maintaining
plant biodiversity in the future.
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