sutherland2012_100 проблем экологии

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FORUM
Identi fi cation of 100 fundamental ecological questions
William J. Sutherland 1, Robert P. Freckleton 2, H. Charles J. Godfray 3,
Steven R. Beissinger 4, Tim Benton 5, Duncan D. Cameron 2, Yohay Carmel 6, David A.
Coomes 7, Tim Coulson 8, Mark C. Emmerson 9, Rosemary S. Hails 10, Graeme C. Hays 11,
Dave J. Hodgson 12, Michael J. Hutchings 13, David Johnson 14, Julia P. G. Jones 15,
Matt J. Keeling 16, Hanna Kokko 17, William E. Kunin 18, Xavier Lambin 14, Owen T. Lewis 3,
Yadvinder Malhi 19, Nova Mieszkowska 20, E. J. Milner-Gulland 21, Ken Norris 22,
Albert B. Phillimore 23, Drew W. Purves 24, Jane M. Reid 14, Daniel C. Reuman 21,25 ,
Ken Thompson 2, Justin M. J. Travis 14, Lindsay A. Turnbull 26, David A. Wardle 27 and
Thorsten Wiegand 28
1Department of Zoology, Conservation Science Group, Cambridge University, Downing Street, Cambridge, CB2 3EJ,
UK; 2Department of Animal and Plant Sciences, University of Shef field, Shef field, S10 2TN, UK; 3Department of
Zoology, University of Oxford, South Parks Road, Oxford, OX1 3PS, UK; 4Department of Environmental Science,
Policy & Management, 137 Mulford Hall #3114, University of California, Berkeley, CA, 94710-3114, USA; 5Global
Food Security Programme, University of Leeds, c/o IICB, Leeds, LS2 9JT, UK; 6Faculty of Civil and Environmental
Engineering, Techion-Israel Institute of Technology, Haifa, Israel; 7Forest Conservation and Ecology Group, Dept of
Plant Sciences, University of Cambridge, Downing Street, Cambridge, CB1 3EA, UK; 8Department of Biology, Imperial
College London, Silwood Park Campus, Ascot, Berkshire, SL5 7PY, UK; 9School of Biological Sciences, Queen ’s
University Belfast, 97 Lisburn Road, Belfast, BT9 7BL, Northern Ireland, UK; 10CEH Wallingford, Maclean Building,
Crowmarsh Gifford, Wallingford, Oxon, OX10 8BB, UK; 11Department of Biosciences, College of Science, Swansea
University, Singleton Park, Swansea, SA1 8PP, UK; 12Centre for Ecology and Conservation, College of Life and
Environmental Sciences, University of Exeter, Cornwall Campus, Penryn, Cornwall, TR10 9EZ, UK; 13School of Life
Sciences, University of Sussex, Falmer, Brighton, Sussex, BN1 9QG, UK; 14Institute of Biological and Environmental
Sciences, University of Aberdeen, Aberdeen, AB14 3UU, UK; 15School of Environment, Natural Resources &
Geography, Thoday Building, Bangor University, Bangor, LL57 2UW, UK; 16School of Life Sciences & Maths Institute
University of Warwick, Gibbet Hill Road, Coventry, CV4 7AL, UK; 17Division of Ecology, Evolution and Genetics,
Research School of Biology, Australian National University, Canberra, ACT 0100, Australia; 18School of Biology,
University of Leeds, Leeds, LS1 9JT, UK; 19Environmental Change Institute, School of Geography and the
Environment, University of Oxford, Oxford, OX1 3QY, UK; 20Marine Biological Association of the UK, Citadel Hill,
Plymouth, PL1 2PB, UK; 21Department of Ecology and Evolution, Silwood Park Campus, Imperial College London,
Buckhurst Road, Ascot, Berks, SL5 7PY, UK; 22Centre for Agri-Environmental Research, University of Reading,
Reading, PO Box 137, RG6 6AR, UK; 23Institute of Evolutionary Biology, University of Edinburgh, Edinburgh, EH9
3JT, UK; 24Computational Ecology and Environmental Science group, Microsoft Research, 7 JJ Thomson Ave,
Cambridge, CB3 0FP, UK; 25Rockefeller University, 1130 York Ave, New York, NY, 10065, USA; 26Institute of
Evolutionary Biology and Environmental Studies, University of Zurich, Winterthurerstrasse 190, 8057 Zurich,
Switzerland; 27Department of Forest Ecology and Management, Faculty of Forestry, Swedish University of Agricultural
Sciences, S901-83 Ume  a, Sweden; and 28Department of Ecological Modelling, UFZ Helmholtz Centre for
Environmental Research-UFZ, Permoserstr 15, 04318, Leipzig, Germany
Summary
1. Fundamental ecological research is both intrinsically interesting and provides the basic knowledge
required to answer applied questions of importance to the management of the natural world. The
100th anniversary of the British Ecological Society in 2013 is an opportune moment to re flect on
the current status of ecology as a science and look forward to high-light priorities for future work.
Correspondence author. E-mail: w.sutherland@zoo.cam.ac.uk
© 2013 The Authors. Journal of Ecology ©2013 British Ecological Society
Journal of Ecology 2013, 101 ,58 –67 doi: 10.1111/1365-2745.12025

2.To do this, we identified 100 important questions of fundamental importance in pure ecology.
We elicited questions from ecologists working across a wide range of systems and disciplines. The
754 questions submitted (listed in the online appendix) from 388 participants were narrowed down
to thefinal 100 through a process of discussion, rewording and repeated rounds of voting. This was
done during a two-day workshop and thereafter.
3.The questions reflect many of the important current conceptual and technical pre-occupations of
ecology. For example, many questions concerned the dynamics of environmental change and com-
plex ecosystem interactions, as well as the interaction between ecology and evolution.
4.The questions reveal a dynamic science with novel subfields emerging. For example, a group of
questions was dedicated to disease and micro-organisms and another on human impacts and global
change reflecting the emergence of new subdisciplines that would not have been foreseen a few dec-
ades ago.
5.The list also contained a number of questions that have perplexed ecologists for decades and are
still seen as crucial to answer, such as the link between population dynamics and life-history evolu-
tion.
6.Synthesis. These 100 questions identified reflect the state of ecology today. Using them as an
agenda for further research would lead to a substantial enhancement in understanding of the disci-
pline, with practical relevance for the conservation of biodiversity and ecosystem function.
Key-words:community ecology, ecology, ecosystems, evolutionary ecology, population ecology,
research priorities
Introduction
Ecologists seek to understand how organisms interact with
each other and the abiotic environment, and also apply this
knowledge to the management of populations, communities
and ecosystems, and the services they provide. Ecologists
todayfind it relatively straightforward to list the major
applied challenges facing thefield. Previous exercises in
which applied ecologists or plant scientists have come
together to draw up lists of the most important questions fac-
ing thefield have revealed a diverse, complex and sometimes
daunting set of challenges (Sutherlandet al.2006, 2009,
2010; Griersonet al.2011). Similar exercises, providing a list
of the major unanswered questions in basic ecology, have
rarely been attempted (but see Thompsonet al.2001). This is
not thefirst time that the British Ecological Society has used
an anniversary as a prompt for an exercise of this type. For
its 75th anniversary in 1988, Cherret (1989) identified the
central existing concepts. The aim of the current exercise was
to look forward to identify key issues.
Such an exercise may be used to evaluate the current state
of the discipline and where its challenges lie. It also helps
to identify areas of research that have the potential to
advance the science of ecology significantly. Furthermore, it
may be particularly valuable as a reference line for future
evaluations of progress in ecology. The last two decades
have seen debates on whether general laws in ecology could
be identified (Moffat 1994; Lawton 1999; Ghilarov 2001;
Dodds 2009; Colyvan & Ginzburg 2012) and the extent to
which ecology is making progress (Abrahamson, Whitham
& Price 1989; Belovskyet al.2004; O’Connor 2000;
Graham & Dayton 2002). The current exercise could add a
concrete dimension to these debates by identifying keyissues and providing an agenda against which progress can
be assessed.
The fundamental aim of ecology is to increase understand-
ing of how organisms interact with the biotic and abiotic
environment rather than address a particular societal, conser-
vation or economic problem. We sought to draw up a list of
important questions facing ecology, with an emphasis on fun-
damental science. Participants were therefore asked to rank
questions by how they would advance ecological science,
rather than by the direct importance of the answer to the
major problems facing society and humanity. Our aim is thus
to set an agenda for means of improving our understanding of
fundamental ecology. There was no attempt to build in con-
sideration of possible application in the future: despite an
increase in horizon-scanning activities (e.g. Sutherlandet al.
2008, 2012), it is inherently difficult to predict what science
will eventually be useful.*
Materials and methods
APPROACH
Our aim was to identify 100 important unanswered questions in basic
ecology. We wanted to avoid very broad, general questions and
instead sought those describing a challenge that could be tackled with
the concerted effort of a small group of researchers or perhaps
through a research programme supported by a limited number of
research grants. As summarized in Table 1, we adopted a previously
used methodology (e.g. Sutherlandet al.2011a) as described in detail
*Benjamin Franklin said that asking the worth of a new discovery
was like saying“What is the use of an infant?”. This is not an argu-
ment that all basic science is equally good, but it is an argument that
the best basic science may have unimaginably important applications.
©2013 The Authors. Journal of Ecology©2013 British Ecological Society,Journal of Ecology,101,58–67
Ecological questions59

in Sutherlandet al.(2011b), which places great emphasis on making
the process to identify the most important questions rigorous, demo-
cratic and transparent.
Participants, which included an editor from each of thefive BES
journals, were selected by WJS, RPF and HCJG after broader consulta-
tion to cover a wide range of approaches to ecology. The attendees
were invited based on their track records of publishing significant sci-
ence in international journals, which we hoped demonstrated their
knowledge of the cutting edge of their subjects. For logistical and
financial reasons, the participants were predominately from the UK;
each is an author of this paper. The attendees were encouraged to con-
sult widely resulting in the active participation of 388 people (includ-
ing those who attended preparatory workshops and discussions, or who
responded to emails, but not those who were sent but did not respond
to emails). The 754 questions submitted are listed in Appendix 1.
The questions were initially assigned to 12 broad themes reflecting
areas of ecology defined by subject or methodological approach. Par-
ticipants were asked to identify and vote for the 6–12 most important
questions in those sections they felt competent to comment on and
suggest rewording where appropriate. All participants were sent and
asked to reflect on the results of the voting and the reworded ques-
tions before the meeting.
A two-day workshop was held at the British Ecological Society’s
headquarters at Charles Darwin House, London, in April 2012. Ques-
tions within each of the themes were considered by working groups
(four consecutive rounds of three parallel sessions). Panel chairs identi-
fied duplicate questions (and ensured that duplication did not lead to
dilution of votes for a particular topic), those that had already been
answered, and those that could be improved by further rephrasing. Par-
ticipants were also encouraged to support potentially important ques-
tions that had not attracted many votes if they considered them
overlooked because of their subject area, because they were in subfields
that were out of fashion, or simply because they were poorly expressed.
The chairs moderated a discussion in which questions that were unli-
kely to make thefinal 100 were quickly excluded before a short list of
18 important questions to be taken to the plenary sessions were agreed.
The latter were divided into three sets of six questions ranked‘bronze’,
‘silver’and‘gold’in the order of increasing importance. Chairs were
asked to ensure the process was democratic with all views respected,
and decisions were made by voting conducted as a show of hands.
The second stage of the workshop consisted of two sets of two par-
allel sessions each of which refined the questions from three of the ini-
tial working groups. Participants werefirst asked to examine the 18
(396) gold questions and remove any duplicates, improve the word-
ing where necessary and demote to the silver section any which on fur-
ther discussion were thought to be of less importance. The 18 bronze
questions were then examined to see whether they contained any that
should be elevated to the silver category. Finally, voting took place toidentify the 20 top questions that formed a new gold group incorporat-
ing the existing gold questions and the most highly supported silver
questions. Further discussion and voting chose from the old silver cate-
gory (and sometimes the bronze) sets offive questions that formed
new silver, bronze and a new category of‘nickel’questions.
In afinal plenary session, the 80 (4920) gold questions were con-
sidered in turn with further elimination of duplicates and major over-
laps. Questions which on further consideration were thought not to be
of the highest importance were demoted to silver, with further voting
when there was no clear consensus. Using the same procedures, partici-
pants were then asked to identify whether any of the questions classi-
fied as nickel should be moved into bronze, and then whether those in
the bronze and following that the silver category should be promoted
or demoted. Thefi
nal rounds of voting chose the most important silver
questions to join the gold questions and so make up thefinal 100.
This voting process was devised so that at each stage the previous
decisions were influential but could also be overruled. It also provided
the opportunity to deal with similar questions that came from different
initial parallel sessions. Furthermore, questions from different groups
were compared against each other to ensure that they were of equivalent
importance and to reduce possible artefacts, for example caused by a dis-
proportionate number of questions initially suggested in one subject area.
Following the workshop, an extensive editing process was carried
out which identified some overlooked ambiguities and duplications. A
final email poll was conducted to decide the fate of the last few can-
didates for inclusion.
LIMITATIONS
Any undertaking such as this of course has limitations (Sutherland
et al.2011b). The most important caveat is that the questions posed
and shortlisted are very likely to be influenced by the interests and
expertise of the participants. Efforts were made to solicit questions
and select attendees from across the full breadth of the subject, but
inevitably biases will remain. In total, 388 people contributed ques-
tions, and there were 37 participants in thefinal workshop. The
majority of the participants were from the UK, and hence, there is a
geographical bias, although we did have attendees from continental
Europe, the US, and Australia, and most participants have many col-
laborators and often conductfieldwork around the globe. We also
invited participants with experience in a range of taxa, including
plants, animals and microbes from both aquatic and terrestrial sys-
tems, to reduce possible taxonomic biases.
The initial division into themes may have limited lateral thinking,
and sometimes, it was not clear where questions should best be placed;
the plenary session andfinal editing was designed to address this issue.
As mentioned previously, there was a tendency to pose broad questions
rather than the more focussed question we were aiming for. There is a
tension between posing broad unanswerable questions and those so
narrow that they cease to be perceived as fundamental. A possible
solution to this in a further exercise might be to define sets of specific
or tactical questions nested within overarching strategic questions.
Results
THE QUESTIONS
The questions here are presented by subject, but not in rank
order. For some questions (e.g. 8, 9, 12, 23, 54), there may
already be a good theoretical understanding but empirical sup-
port for the theory is still lacking.
Table 1.The process used for reducing the submitted questions into
thefinal list of 100. Thefirst stage involved prioritizing the complete
set of questions. Each subsequent stage used the ranking of the previ-
ous stage to influence the narrowing of the list
1. 754 questions categorized into 12 groups and ranked by voting
before the meeting.
2. Twelve sessions, each dealing with one group, identify 6 highest
priority‘gold’questions, 6‘silver’and 6‘bronze’.
3. Four sessions, each taking output from three sessions
in stage (2), identifying 20‘gold’questions, 5‘silver’,
5‘bronze’and 5‘nickel’.
4. Plenary session identifying the top 100.
©2013 The Authors. Journal of Ecology©2013 British Ecological Society,Journal of Ecology,101,58–67
60W. J. Sutherlandet al.

EVOLUTION AND ECOLOGY
Ecology and evolution share a broad interface, with both
fields recognizing the value of an inter-disciplinary perspec-
tive. Interest in the role of abiotic conditions and biotic inter-
actions as drivers of natural selection (Questions 1, 3) is long
standing (Darwin 1859) and remains an active area of
research (Kingsolveret al.2012). More recent, in light of evi-
dence for very rapid evolution is a focus on eco-evolutionary
dynamics (Schoener 2011). Population dynamic can influence
selection from one generation to the next, but at the same,
time life history may evolve and feedback upon population
dynamics. This research programme is dissolving the distinc-
tion between evolutionary and ecological time-scales and is
represented by several of the questions in this section that
address aspects of the interplay between life-history evolution
and population dynamics (5, 6, 8). Despite calls for ecologists
to engage with the emergentfield of epigenetics (Bossdorf,
Richards & Pigliucci 2008), it is represented by a solitary
question (4), the breadth of which highlights how just little is
known from either a theoretical or empirical perspective.
Reflecting some of the range of influences that evolution has
on ecology, andvice versa, questions with an explicit evolu-
tionary component also appear under the Populations (11,
14), Communities and Diversity (48, 56) and Human impacts
and global change (74, 82) sections.
1What are the evolutionary consequences of species
becoming less connected through fragmentation or more
connected through globalization?
2To what extent can evolution change the scaling rela-
tionships that we see in nature?
3How local is adaptation?
4What are the ecological causes and consequences of epi-
genetic variation?
5What are the relative contributions of different levels of
selection (gene, individual, group) to life-history evolu-
tion and the resulting population dynamics?
6What selective forces cause sex differences in life his-
tory and what are their consequences for population
dynamics?
7How should evolutionary and ecological theory be mod-
ified for organisms where the concepts of individual and
fitness are not easily defined (e.g. fungi)?
8How do the strength and form of density dependence
influence feedbacks between population dynamics and
life-history evolution?
9How does phenotypic plasticity influence evolutionary
trajectories?
10What are the physiological bases of life-history trade-
offs?
POPULATIONS
Understanding and predicting the spatio-temporal dynamics of
populations remains a central goal in ecology (Davidson &
Andrewartha 1948; Hanski 1998; Alexanderet al.2012). This
requires detailed understanding of how demographic ratesvary and covary through space and time as well as the under-
lying causes. Several questions reflect the drive to gain this
understanding (e.g.17, 18, 23). The recent accumulation of
evidence suggesting that evolutionary processes can occur
rapidly enough to influence population dynamics at a range of
spatial scales has resulted in renewed emphasis on joint anal-
ysis of population dynamics and life-history evolution (Pelle-
tier, Garant & Hendry 2009; Schoener 2011), which is
reflected in questions 20, 23). Dispersal is a key process
determining spatial population dynamics and technological
innovations have revolutionized our ability to measure indi-
vidual movement trajectories (Cagnacciet al.2010). Under-
standing the causes of variability in dispersal and their
consequences for spatial dynamics across different spatial
scales remains a major focus of ecological enquiry and future
major challenges are emphasized in several of the questions
(13–16). While we surmise that processes operating atfine
spatial and/or temporal scales are likely to impact dynamics
at large spatial scales such as species’ranges, there remains
an urgent need for new methods that enable us to link local
processes to large-scale spatial dynamics (12) (e.g. Helmuth
et al.2006). This linkage will help our understanding of how
local population dynamics link to macroecological patterns
and dynamics (11, 19), as well as improve predictions of pop-
ulation dynamics.
11What are the evolutionary and ecological mechanisms
that govern species’range margins?
12How can we upscale detailed processes at the level of
individuals into patterns at the population scale?
13How do species and population traits and landscape
configuration interact to determine realized dispersal
distances?
14What is the heritability/genetic basis of dispersal and
movement behaviour?
15Do individuals in the tails of dispersal or dormancy
distributions have distinctive genotypes or phenotypes?
16How do organisms make movement decisions in rela-
tion to dispersal, migration, foraging or mate search?
17Do different demographic rates vary predictably over
different spatial scales, and how do they then combine
to influence spatio-temporal population dynamics?
18How does demographic and spatial structure modify
the effects of environmental stochasticity on population
dynamics?
19How does environmental stochasticity and environmen-
tal change interact with density dependence to generate
population dynamics and species distributions?
20To what degree do trans-generational effects on life
histories, such as maternal effects, impact on popula-
tion dynamics?
21What are the magnitudes and durations of carry-over
effects of previous environmental experiences on an
individual’s subsequent life history and consequent
population dynamics?
22What causes massive variability in recruitment in some
marine systems?
©2013 The Authors. Journal of Ecology©2013 British Ecological Society,Journal of Ecology,101,58–67
Ecological questions61

23How does covariance among life-history traits affect
their contributions to population dynamics?
24What is the relative importance of direct (consumption,
competition) vs. indirect (induced behavioural change)
interactions in determining the effect of one species on
others?
25How important is individual variation to population,
community and ecosystem dynamics?
26What demographic traits determine the resilience of
natural populations to disturbance and perturbation?
DISEASE AND MICRO-ORGANISMS
While the study of infectious disease is often seen as a branch
of medical science, the way that all micro-organisms (from
parasites to commensalists to mutualists) interact with their
hosts and their environment clearlyfits within the remit of
ecology. Indeed, for many years, the study of infectious dis-
eases (e.g. Anderson & May 1992) has used ecological con-
cepts to improve our understanding of public-health issues.
Furthering understanding of the regulation of disease continues
to require knowledge of basic microbiology and there is grow-
ing realization within the discipline of ecology that the abun-
dance, diversity and function of micro-organisms have
fundamental roles in shaping ecosystems. This view appears to
be borne-out by the selected questions, which tend to focus on
interactions between micro-organisms and larger organisms (e.
g. 28–31). The rapid development and application of molecular
techniques continues to reveal a previously hidden diversity of
micro-organisms, particularly in complex environments such
as soils (Roslinget al.2011). Population genomics has pro-
vided insight into the genetic mechanisms using which micro-
organisms interact with, and help shape, their environment (e.
g. Martinet al.2008, 2010), and this calls for a better under-
standing of the importance of microbial genotypic diversity for
ecosystems (Johnsonet al.2012; 29, 30). The questions also
reveal the need to test the suitability of general ecological the-
ory to microbial systems (35), and to determine how experi-
mental microbial systems can inform and develop ecological
theory (36) that has often been derived from or applied to ma-
croorganisms (Prosseret al.2007).
27How important are multiple infections in driving dis-
ease dynamics?
28What is the role of parasites and mutualists in gener-
ating and maintaining host species diversity?
29How does below-ground biodiversity affect above-
ground biodiversity, andvice versa?
30What is the relationship between microbial diversity
(functional type, species, genotype) and community
and ecosystem functioning?
31To what extent is macroorganism community compo-
sition and diversity determined by interactions with
micro-organisms?
32What is the relative importance of biotic vs. abiotic
feedbacks between plants and soil for influencing
plant growth?33How do symbioses between micro-organisms and
their hosts influence interactions with consumers and
higher trophic levels?
34In what ecological settings are parasites key regula-
tors of population dynamics?
35Do the same macroecological patterns apply to
micro-organisms and macroorganisms, and are they
caused by the same processes?
36What can we learn from model communities of
micro-organisms about communities of macroorgan-
isms?
37How does intraspecific diversity contribute to the
dynamics of host-parasite and mutualistic interactions?
COMMUNITIES AND DIVERSITY
Some of the most challenging questions in ecology concern
communities: sets of co-occurring species. For much of the last
century, ecologists have typically interpreted the diversity and
composition of communities as the outcome of local-scale pro-
cesses, both biotic (e.g. competition and predation) and abiotic
(e.g. temperature and nutrients). However, some have chal-
lenged this view, and emphasize the importance of chance (e.g.
Hubbell 2001) and large-scale biogeography and evolutionary
history (e.g. Ricklefs 2008) and many issues remain (e.g. 47,
48, 50, 52). Ecologists need to resolve the extent to which the
structure and dynamics of ecological communities can be pre-
dicted from the traits of their component species (38–40).
Understanding the nature and ramifications of the networks of
interactions among species remains a major priority (e.g. 41,
42), as does understanding the role of environmental variability
through space and time (39, 43, 45). A developing area of
emphasis–interfacing with questions listed under the‘ecosys-
tems’heading–is on the functioning of ecological communities
in relationship to their diversity, composition and structure. A
large body of experimental research has explored these rela-
tionships, but most experiments are necessarily restricted to
small sets of species, often drawn from a single trophic level.
Many important questions about the attributes of‘real’ecologi-
cal communities in relation to their functioning remain unan-
swered (e.g. 39, 44, 49).
38How can we use species’traits as proxies to predict
trophic interaction strength?
39How well can community properties and responses to
environmental change be predicted from the distribu-
tion of simple synoptic traits, e.g. body size, leaf area?
40How do species traits influence ecological network
structure?
41When, if ever, can the combined effect of many weak
interactions, which are difficult to measure, be greater
than the few strong ones we can easily measure?
42How widespread and important are indirect interactions
(e.g. apparent competition, apparent mutualism) in eco-
logical communities?
43How do spatial and temporal environmental heteroge-
neity influence diversity at different scales?
©2013 The Authors. Journal of Ecology©2013 British Ecological Society,Journal of Ecology,101,58–67
62W. J. Sutherlandet al.

44How does species loss affect the extinction risk of the
remaining species?
45What is the relative importance of stochastic vs. deter-
ministic processes in controlling diversity and composi-
tion of communities, and how does this vary across
ecosystem types?
46How do we predict mechanistically how many species
can coexist in a given area?
47To what extent is local species composition and diver-
sity controlled by dispersal limitation and the regional
species pool?
48What are the contributions of biogeographical factors
and evolutionary history in determining present day
ecological processes?
49To what extent is primary producer diversity a driver
of wider community diversity?
50How relevant are assembly rules in a world of biologi-
cal invasion?
51What is the relative importance of trophic and non-
trophic interactions in determining the composition of
communities?
52How important are dynamical extinction-recolonization
equilibria to the persistence of species assemblages in
fragmented landscapes?
53Which mechanisms allow the long-term coexistence of
grasses and woody plants over a wide range of ecosys-
tems?
54How do resource pulses affect resource use and inter-
actions between organisms?
55How important are rare species in the functioning of
ecological communities?
56What is the feedback between diversity and diversifi-
cation?
57What are the functional consequences of allelopathy
for natural plant communities?
ECOSYSTEMS AND FUNCTIONING
Our understanding of how biotic and abiotic factors drive the
functioning of ecosystems has advanced rapidly over the last
two decades, in part as a consequence of a growing degree of
integration of community-level and ecosystem-level ecology.
As such, we now have a much better understanding of how
community diversity and composition influence ecosystem pro-
cesses, the resistance and resilience of ecosystems to environ-
mental perturbations, and feedbacks between the producer and
decomposer components of ecosystems. There is also growing
awareness of how ecosystems respond to global environmental
changes, their capacity to regulatefluxes of carbon and nutri-
ents, and their interactions with the Earth climate system, but
challenges remain (e.g. 61, 69, 72). Future challenges for eco-
system science, as reflected in the questions, include being able
to make predictions about ecosystems undergoing catastrophic
transitions (e.g. 58–60, 71) (Schefferet al.2009), better under-
standing the role of spatial scale in driving ecosystem processes
(e.g. 63), and extending our rapidly growing knowledge of eco-
logical networks (Bascompte 2009) to study the functioning ofecosystems (e.g. 65). Another major challenge is to better
understand the responses of ecosystems to realistic scenarios of
biodiversity change through the simultaneous processes of
extinction (Cardinaleet al.2012) and invasion (Simberloff
et al.2012) (e.g. 61–63, 68).
58Which ecosystems are susceptible to showing tipping
points and why?
59How can we tell when an ecosystem is near a tipping
point?
60Which factors and mechanisms determine the resil-
ience of ecosystems to external perturbations and
how do we measure resilience?
61Which ecosystems and what properties are most sen-
sitive to changes in community composition?
62How is ecosystem function altered under realistic sce-
narios of biodiversity change?
63What is the relative contribution of biodiversity at
different levels of organization (genes, species rich-
ness, species identity, functional identity, functional
diversity) to ecosystem functioning?
64What are the generalities in ecosystem properties and
dynamics between marine, freshwater and terrestrial
biomes?
65How does the structure of ecological interaction net-
works affect ecosystem functioning and stability?
66How does spatial structure influence ecosystem func-
tion and how do we integrate within and between
spatial scales to assess function?
67How do nutrients other than nitrogen and phosphorus
(and iron in the sea) affect productivity in ecosys-
tems?
68To what extent is biotic invasion and native species
loss creating ecosystems with altered properties?
69Are there globally significant ecosystem functions
provided by poorly known ecosystems (e.g. deep
oceans, ground water)?
70Which, if any, species are functionally redundant in
the context of stochastic or directional environmental
changes?
71Is hysteresis the exception or the norm in ecological
systems?
72Can we predict the responses of ecosystems to envi-
ronmental change based on the traits of species?
HUMAN IMPACTS AND GLOBAL CHANGE
It is increasingly recognized that current ecological dynam-
ics and ecosystem function occurs within the context of a
human-dominated planet (Marris 2011) and that many eco-
systems have been altered and affected by humans since
pre-history (Gillet al.2009; Doughty, Wolf & Field 2010).
Human impacts on ecosystems include direct impacts on
habitats such as land conversion andfire use, habitat modi-
fication (such as selective logging or changing in drainage
of wetlands), changes in connectivity (fragmentation or
globalization) as well as changes in species composition
through removal (due to harvesting or pest control) or intro-
©2013 The Authors. Journal of Ecology©2013 British Ecological Society,Journal of Ecology,101,58–67
Ecological questions63

duction (accidental or otherwise) of species. These impacts
generate many important questions (73–75, 85, 86, 88, 89).
Another suite of human impacts is more indirect but per-
haps even more pervasive; through our alteration of the cli-
mate (both its mean state and variability; IPCC 2007;
Hannah 2011) and changes in the biogeochemistry of the
atmosphere and oceans (Heimann & Reichstein 2008; Do-
neyet al.2009). These alterations raise questions about
what determines how and how fast particular species
respond to such change (82, 83), how communities of spe-
cies interact and respond to change (80, 81, 87), and
whether past rates of change can yield insights into likely
ecological responses to current and future change (84).
Another set of global change ecology questions is centred
on how the functioning of the biosphere as a whole is
affected by global change, and what role the biosphere
plays in the response of the atmosphere to human impacts,
through the carbon and water cycles and other major bio-
geochemical cycles (76–79).
73What is the magnitude of the‘extinction debt’fol-
lowing the loss and fragmentation of natural habitats,
and when will it be paid?
74What is the role of evolution in recovery from
exploitation and responses to other forms of relaxed
selection?
75What are the indirect effects of harvesting on ecosys-
tem structure and dynamics?
76What are the major feedbacks and interactions
between the Earth’s ecosystems and the atmosphere
under a changing climate?
77What are the key determinants of the future magni-
tude of marine and terrestrial carbon sinks?
78How will atmospheric change affect primary produc-
tion of terrestrial ecosystems?
79How will ocean acidification influence primary pro-
duction of marine ecosystems?
80To what extent will climate change uncouple trophic
links due to phenological change?
81How do natural communities respond to increased
frequencies of extreme weather events predicted
under global climate change?
82In the face of rapid environmental change, what
determines whether species adapt, shift their ranges
or go extinct?
83What determines the rate at which species distribu-
tions respond to climate change?
84To what extent can we extrapolate from palaeoeco-
logical range shifts to understand 21st-century
change?
85Under what circumstances do landscape structures
such as corridors and stepping stones play important
roles in the distribution and abundance of species?
86To what extent will the breakdown of biogeographi-
cal barriers (e.g. the more permanent opening of the
Northwest Passage) lead to sustained changes in local
diversity?87How do interspecific interactions affect species
responses to global change?
88What are the ecosystem impacts of world-wide top
predator declines?
89What is the legacy of Pleistocene megafauna extinc-
tions on contemporary ecosystems?
METHODS
Over the past two decades, the practice of ecology has been revo-
lutionized by the development of new technologies, and further
developments will continue to be an important stimulus to new
research. Important advances include the increase in the avail-
ability and speed of computers, new molecular approaches for
resolving diversity and dispersal, barcoding techniques that per-
mit rapid identification and even phylogeny building at the com-
munity level, the development of new statistical methods (e.g.
mixed models and Bayesian statistics, e.g. Bolkeret al.2009),
monitoring tools such as remote sensing (Asneret al.2008) and
geo-tagging of individuals (Blocket al.2001). There is also
increasing use of citizen science to conduct ecological and evo-
lutionary studies (e.g. Worthingtonet al.2012).This set of ques-
tions reflects on the methods used to conduct research in ecology
and the lessons that can be drawn from previous ecological stud-
ies, for example whether previous predictions have been suc-
cessful or erroneous (91, 92, 94). It encompasses new
technology (95, 96), as well as the development of new tools and
inter-disciplinary links (90, 99, 100). The development of new
tools for measuring and monitoring is an important focus (96,
98), and this includes developing methods to model the observa-
tion process itself (99).
90What unexploited theories used by other disciplines
could inform ecology, and vice versa?
91How do we best develop and exploit empirical
model systems for understanding natural systems?
92How successful have past ecological predictions
been and why?
93What is the nature of published ecological errors
and how do errors affect academic understanding
and policy?
94How is our understanding of ecology influenced by
publication bias?
95What new technologies would most advance ecolog-
ical understanding?
96How do we combine multiple scales and types of
monitoring (fromfield to earth observation) to make
robust ecological inferences?
97To what extent are widely studied ecological pat-
terns (species-abundance distribution, species-area
relationship, etc.) the outcomes of statistical rather
than ecological processes?
98What are the most appropriate baselines for deter-
mining the magnitude and direction of ecological
changes?
99How much does modelling feedbacks from the
observation process, such as the responses of organ-
©2013 The Authors. Journal of Ecology©2013 British Ecological Society,Journal of Ecology,101,58–67
64W. J. Sutherlandet al.

isms to data collectors, improve our ability to infer
ecological processes?
100How can the feedbacks between human behaviour
and ecological dynamics be accounted for in ecolog-
ical models?
Discussion
KNOWLEDGE GAPS IN ECOLOGY
Collaborative projects to highlight and prioritize unanswered
research questions allow researchers to review and reflect on
the current state of a discipline, and how it is likely to
develop in the future. Our list of 100 unanswered questions
includes many that address the nature of fundamental con-
cepts and principles in ecology. For example, some questions
reveal profound knowledge gaps regarding the central mecha-
nisms driving ecosystems [61, 63, 64, 75, 76, 77], communi-
ties [42, 45, 47, 48, 51], and even population dynamics
[11, 19].
All vibrantfields of science have unanswered questions,
but are there characteristics of ecology as a discipline that
might explain why some large knowledge gaps remain after
100 years of intensive research? One explanation of barriers
to progress in ecology maintains that it is a science of
middle numbers (Allen & Hoekstra 1992). In small-number
systems like the solar system, the relationships between the
components, and the state of the system, can often be ade-
quately described by a simple set of equations. In contrast,
in large-number systems such as chemical interactions in
fluids, the behaviour of the system can usually be ade-
quately described using statistical averages because of the
large number of components and the simple nature of their
interactions. Ecological systems unfortunately belong to the
study of middle numbers: they are too complex to describe indi-
vidually, yet their components are too few and their interactions
too complex to be described by statistical dynamics. Com-
pounding this problem is the long time-scale of ecological
dynamics: many interesting phenomena, especially those
involving ecosystems, have decadal time-scales making their
study difficult and leading to a lack of long-term data. Although
great progress has undoubtedly been made in the last 100 years,
we must continue the task of observing, experimenting and
modelling, anticipating the expected, and unexpected, steady
progress and great leaps forward which will result. It would be
interesting to repeat this exercise in 10 or 15 years’time to
monitor progress.
Ecology has its origins in natural history and early publica-
tions tended to be very descriptive and site-specific. Modern
ecology has progressed through the incorporation of highly
sophisticated numerical methods, as well as becoming under-
pinned by a set of strong theories. Some of the questions
identified here are moderately well understood from a theoret-
ical perspective but require more empirical research. It is
instructive to note that volume 1, issue 1 ofJournal of
Ecology, the oldest ecological journal, contained only a single
paper that referenced statistics (Smith 1913) and no paper inthatfirst issue of the journal tested a hypothesis. Modern
ecology is a hugely collaborative discipline. Many of the
questions listed here link to other disciplines within biology
including genetics, epidemiology and evolutionary biology.
Furthermore, while for clarity we have organized the ques-
tions into themes, it is notable that many of the unanswered
questions cut across these rather arbitrary divisions.
FUTURE DIRECTIONS
There have been intermittent calls over the decades for the
development of a general theory of ecology. The desirability
and feasibility of this has been debated extensively (Scheiner
& Willig 2005; Roughgarden 2009). We would agree with
Loreau (2010) that the way forward is not a single monolithic
theory, but increasing the process of merging-related disci-
plines to generate new principles, perspectives, and questions
at the interfaces, thus contributing to the emergence of a new
ecological synthesis transcending traditional boundaries. The
range of questions presented here reflects the diversity of
modern ecology. There is a balance of questions best
answered by theoretical approaches, experiment and observa-
tion and all these approaches will continue to be important.
Global environmental change provides an important context
for current ecological research. Much past ecological theory
was derived for systems thatfluctuated very little around an
average state, but global change is leading to both long-term
shifts in average conditions as well as potentially dramatic
changes in environmental variation. Many of the questions
identified are concerned with understanding how systems will
respond to such changes.
It is encouraging that there was a general consensus that
some areas viewed as hot topics over the last few decades
did not need to be included in our list; evidence that the
discipline is progressing. It was clear from discussions that
questions once considered important had not been defini-
tively answered; but rather that the focus had shifted in the
light of improved understanding and experience. If this
exercise had been conducted 40 years ago then many of
the questions would have involved density dependence and
whether or not it was present in thefield. Today there is a
consensus that density dependence is pervasive, but also
that it may take very different forms and sometimes be
very hard to detect. Looking back, much of the heat of the
discussion involved people misunderstanding each other.
Some questions set 25 years ago would have involved the
search for dynamical deterministic chaos. We now know
that intrinsic and extrinsic (stochastic) forces act together to
determine observed dynamics and looking for pure deter-
ministic chaos has little meaning (in as much as weather
affects population dynamics all species have chaotic
dynamics).
In communities and ecosystems, questions of community
equilibria have evolved into questions about resilience and per-
turbation of communities, or indeed whole ecosystems, and such
thinking has been incorporated in the study of phylogenetic
diversity patterns through time (e.g. Rabosky & Glor 2010).
©2013 The Authors. Journal of Ecology©2013 British Ecological Society,Journal of Ecology,101,58–67
Ecological questions65

CONCLUDING REMARKS
Both the science of ecology and the British Ecological Soci-
ety have come a long way over the last 100 years. In 1913,
the BES was made up of a relatively small group of mostly
British scientists with a focus on studying natural history in
natural environments. Today, it is a dynamic international
organization with members representing academia, industry,
education and NGOs, and coming from more than 80 coun-
tries. These members conduct pure research, answer applied
questions concerning restoration and management, and influ-
ence government policy. They work in protected areas as well
as farmland, post-industrial landscapes and the urban environ-
ment. Despite expanding its initial remit and reaching out far
beyond its membership, the science of ecology remains at the
heart of the BES. In this paper, a large group of ecologists
have prioritized 100 questions they think are the most impor-
tant remaining questions for the science of ecology to tackle.
We do not claim this list to be definitive but hope that it stim-
ulates discussion and exciting new research.
Acknowledgements
We thank the 388 people who participated in workshops and conversations that
resulted in the initial list of questions. The British Ecological Society provided
funding, hosting and, through Heather Mewton and Olivia Hunter, organized
the workshop. Stephanie Prior played a major role in collating questions and
collating materials for the manuscript. Holly Barclay, Yangchen Lin and Jessica
Walsh edited the questions on laptops during the workshop. Andrew Becker-
man, Mike Begon, Alastair Fitter, Kathy Willis and Ken Wilson contributed
questions and scores but were unable to attend the workshop. Bill Bewes pro-
vided the data on distribution of BES members. We thank David Gibson, Mark
Rees and an anonymous referee for their useful comments.
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Received 12 June 2012; accepted 22 October 2012
Handling Editor: David Gibson
Supporting Information
Additional Supporting Information may be found in the online ver-
sion of this article:
Appendix S1.A list of the 754 submitted questions.
©2013 The Authors. Journal of Ecology©2013 British Ecological Society,Journal of Ecology,101,58–67
Ecological questions67