Austral Ecology (2012) ••, ••–••

Value of long-term ecological studies aec_2351 1..13

DAVID B. LINDENMAYER,1* GENE E. LIKENS,2,3 ALAN ANDERSEN,4

DAVID BOWMAN,7 C. MICHAEL BULL,8 EMMA BURNS,1 CHRIS R. DICKMAN,12,13
ARY A. HOFFMANN,16,17 DAVID A. KEITH,14,15 MICHAEL J. LIDDELL,18
ANDREW J. LOWE,9,10,11 DANIEL J. METCALFE,19 STUART R. PHINN,20,21
JEREMY RUSSELL-SMITH,5,6 NIKKI THURGATE9,10 AND GLENDA M. WARDLE12,13
1
Fenner School of Environment and Society, The Australian National University, Canberra, ACT 0200,
Australia (Email: david.lindenmayer@anu.edu.au); 2Cary Institute of Ecosystem Studies, Millbrook,
USA; 3Department of Limnology, Uppsala University, Uppsala, Sweden; 4CSIRO Ecosystem Sciences,
Tropical Ecosystems Research Centre, 5Bushfires NT and 6Research Institute for the Environment and
Livelihoods, Charles Darwin University, Darwin, Northern Territory, 7School of Plant Science, The
University of Tasmania, Hobart, Tasmania, 8School of Biological Sciences, Flinders University,
9
Australian Centre for Evolutionary Biology and Biodiversity and 10School of Earth and Environmental
Sciences, University of Adelaide, 11State Herbarium of South Australia, Science Resource Centre,
Department of Environment and Natural Resources, Adelaide, South Australia, 12Desert Ecology
Research Group and 13Institute of Wildlife Research, School of Biological Sciences, The University of
Sydney, 14School of Biological, Earth and Environmental Sciences, University of New South Wales,
15
NSW National Parks & Wildlife Service, Sydney, New South Wales, Departments of 16Zoology and
17
Genetics, Bio21 Institute, The University of Melbourne, Melbourne,Victoria, 18School of Pharmacy
and Molecular Sciences, James Cook University, Cairns, 19CSIRO Ecosystem Sciences, Tropical Forest
Research Centre, Atherton, 20Terrestrial Ecosystem Research Network and 21School of Geography,
Planning and Environmental Management, The University of Queensland, Brisbane, Queensland,
Australia

Abstract Long-term ecological studies are critical for providing key insights in ecology, environmental change,
natural resource management and biodiversity conservation. In this paper, we briefly discuss five key values of such
studies. These are: (1) quantifying ecological responses to drivers of ecosystem change; (2) understanding complex
ecosystem processes that occur over prolonged periods; (3) providing core ecological data that may be used to
develop theoretical ecological models and to parameterize and validate simulation models; (4) acting as platforms
for collaborative studies, thus promoting multidisciplinary research; and (5) providing data and understanding at
scales relevant to management, and hence critically supporting evidence-based policy, decision making and the
management of ecosystems. We suggest that the ecological research community needs to put higher priority on
communicating the benefits of long-term ecological studies to resource managers, policy makers and the general
public. Long-term research will be especially important for tackling large-scale emerging problems confronting
humanity such as resource management for a rapidly increasing human population, mass species extinction, and
climate change detection, mitigation and adaptation. While some ecologically relevant, long-term data sets are now
becoming more generally available, these are exceptions. This deficiency occurs because ecological studies can be
difficult to maintain for long periods as they exceed the length of government administrations and funding cycles.
We argue that the ecological research community will need to coordinate ongoing efforts in an open and
collaborative way, to ensure that discoverable long-term ecological studies do not become a long-term deficiency.
It is important to maintain publishing outlets for empirical field-based ecology, while simultaneously developing
new systems of recognition that reward ecologists for the use and collaborative sharing of their long-term data sets.
Funding schemes must be re-crafted to emphasize collaborative partnerships between field-based ecologists,
theoreticians and modellers, and to provide financial support that is committed over commensurate time frames.

Key words: ecological change, ecological theory, long-term ecological study, modelling, vulnerability assessment.

*Corresponding author.
Accepted for publication December 2011.

© 2012 The Authors doi:10.1111/j.1442-9993.2011.02351.x

Journal compilation © 2012 Ecological Society of Australia
2 D. B. L I N D E N M AY E R ET AL.
INTRODUCTION
Effective ecosystem management requires a robust
understanding of the long-term dynamics of ecosys-
tems, communities and populations. High-quality
ecological information collected over long periods
can yield valuable insights into changes in ecosystem
structure, key ecological processes and the services
provided by ecosystems (e.g. Odum 1959; Westoby
1991; Daily 1997; Holmes & Sherry 2001; Krebs et al.
2001; Knowlton & Jones 2006; Willis et al. 2008;
Lindenmayer & Likens 2010; Holmes 2011). Indeed,
long-term ecological studies are especially important
for quantifying the complex, multiple, simultaneous,
non-linear interactions that are prevalent in many eco-
systems (Runyoro et al. 1995; Levin 2009; Estes et al.
2011). Some long-term data sets (consisting of field
surveys, instrumental records and maps) are now
becoming available in Australia and other countries
such as the USA (e.g. RAINFOR, NEON, NCEAS,
Fluxnet and Landsat Archives), South Africa (e.g.
SAEON), China (CFERN) and members of the Euro-
pean Union (e.g. LTER-Europe, Exper-ER), and via
the International Long Term Ecological Research
(ILTER). However, there are very few institutionalized
and openly accessible long-term studies; most are ad
hoc, fragmented and have a tenuous existence. Storage
of these data in association with publications is being
encouraged in new journal publication requirements
by some ecological journals, such as those of the
Ecological Society of America.
Our primary aim in this paper is to outline some of
the main values of long-term ecological studies. We
suggest this is important because resource managers,
policy makers and the general public may be unaware of
these values and of the critical role of long-term eco-
logical studies in tackling emerging problems of major
social concern. These include developing ecologically
sustainable resource management strategies in the face
of a rapidly increasing human population, preventing
mass species extinction, and detecting, mitigating and
adapting to rapid climate change. Conversely, without
an explicit recognition of the multiple values provided
by long-term ecological studies and their data sets, we
argue that in the future, important ecological discover-
ies will be overlooked, robust modelling, meta-analyses
and systematic reviews will be difficult or problematic
(Lindenmayer et al. 2010; Lindenmayer & Likens
2011) and, ultimately, evidence-based environmental
policy and management (e.g. Sutherland et al. 2004;
Collaboration for Environmental Evidence 2011) will
be difficult to achieve.Therefore, in the final part of this
paper, we suggest some ways to improve the chances
that existing long-term studies might be maintained
and new ones initiated both in Australia and elsewhere
around the world. The primary geographical focus of
this paper is Australia, but we have included examples
doi:10.1111/j.1442-9993.2011.02351.x
Journal compilation © 2012 Ecological Society of Australia
from other countries to illustrate the values of long-
term ecological studies more broadly. Our intended
audience is the research community, resource manag-
ers and policy makers in natural resource management
and conservation management.
WHAT ARE LONG-TERM ECOLOGICAL
STUDIES?
There are many nuances to discussions about what
constitutes ‘long term’ in ecological research. Strayer
et al. (1986) noted that some workers consider long-
term studies to be those that continue beyond the
generation time of dominant organisms in an ecosys-
tem, or sufficiently long to quantify the key processes
that structure the ecosystem under investigation.
Under this definition, month-long studies of bacterial
assemblages with very rapid generation times could be
considered long-term. Conversely, a 300-year study of
stands of trees in which the dominant trees may live for
500–1000 years would not qualify as long-term. These
considerations are important because they emphasize
the role of variable life spans of different organisms,
but they are not feasible to use for many ecosystem
analyses.
For the purposes of this paper, we use a practical,
operational definition and consider long-term ecologi-
cal studies to be those that systematically and regularly
collect field data from a particular site or set of sites for
more than 10 years.The justification for this somewhat
arbitrary time component is that it provides repeated
sampling of annual growth cycles and it includes
some longer-term bioclimatic cycles that operate over
decadal periods.
It is also important to emphasize what is not long-
term ecological research. What is distinctive is the
number of (continuous) years of the study rather than
any processes that may operate over long time periods.
We therefore do not consider simulation modelling per
se to be long-term work, even though a simulation
model can rather easily make projections for thousands
of years into the future.We also do not consider studies
that substitute space-for-time (e.g. snapshot investiga-
tions; Diamond 1986; Likens 1989a) or retrospective
investigations (sensu Likens 1989a), such as those of
pollen cores, tree rings, otoliths and bio-accumulated
signatures in tissues, to be long-term work per se,
although they can provide long-term perspectives. We
also do not consider haphazard revisits to a site after a
prolonged absence (e.g. Currie & Parry 1999; Smith
et al. 2007) to be programmes of long-term work,
unless the design of the study is focused on responses to
irregular events such as recurring floods, fires or storms.
These efforts, while valuable, do not face the same
challenges that distinguish longitudinal, on-the-
ground, ecological field studies conducted systemati-
cally over many years.
© 2012 The Authors
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THE VALUES AND BENEFITS OF
LONG-TERM ECOLOGICAL DATA SETS
Numerous scientific articles, books, reports, manage-
ment strategies and other documents stress that
long-term ecological studies are needed to answer
important questions about ecosystem, community and
population dynamics. A common aim has been to
identify key ecological processes that give rise to
spatial and temporal patterns in the distribution, abun-
dance or diversity of biota (e.g. Holmes & Sherry
2001; Krebs et al. 2001) or the causal factors under-
pinning environmental problems (Lindenmayer &
Likens 2010). Thus, long-term studies can provide an
improved understanding of complicated ecological
systems (e.g. Strayer et al. 1986; Likens 1989a, 1992;
Goldsmith 1991; Westoby 1991; Spellerberg 1994;
Condit 1995; Likens & Bormann 1995; Thompson
et al. 1998; Franklin et al. 1999; Hubbell 2002; Chave
et al. 2003; Lovett et al. 2007; Krebs et al. 2008;
Dickman et al. 2010). Implicit in our following discus-
sion of the key values of long-term ecological data sets
is that they provide an enduring resource for current
and future ecologists, land managers and policy
makers. In the remainder of this paper, we consider
five key values of long-term ecological studies.
Quantifying ecological responses to
environmental change
Long-term ecological data are often critical for quan-
tifying ecological responses to environmental change
such as natural, human or experimental disturbance
(Likens 1985; Carpenter et al. 1995). For example,
long-term experimental work in southern New South
Wales, Australia has demonstrated how changes in
the landscape context of patches of remnant native
woodland, driven by the establishment and subse-
quent maturation of stands of exotic Pinus radiata
surrounding the patches, can significantly alter
assemblages of birds and other vertebrates (Linden-
mayer et al. 2008) (Fig. 1). The results of this work
have significant implications for the design of exotic
tree plantations, particularly where such land uses are
planned around endangered woodland vegetation
communities (Lindenmayer 2009b). Long-term
studies have uncovered complex shifting top–down
and bottom–up ecological processes on assemblages
of mammals in Australian deserts (Letnic et al. 2011)
as well as other desert ecosystems around the world
(Meserve et al. 2011). These insights have been criti-
cal for informing the management of populations of
native mammals, particularly where there are also
populations of exotic herbivores and predators
(Letnic et al. 2011).
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Journal compilation © 2012 Ecological Society of Australia
3
Long-term studies can indicate the vulnerability of
particular species to environmental changes. Surveys
of flowering time in plant species from the Victorian
high country in Australia have shown marked inter-
specific differences in phenological responsiveness to
changing environmental conditions (Hoffmann et al.
2010). Similarly, Willis et al. (2008, 2010) explored a
150-year data set of plants from Concord, Massachu-
setts, USA and found that some species, including
invasive taxa, changed their phenological traits to
track changes in seasonal timing and increased in
abundance as climatic conditions warmed, while
others, including many native species, flowered at a
fixed time and had become locally extinct. A 23-year
data set from the shortgrass steppe of the Central
Plains of the USA showed decreasing net primary
production by the dominant C4 grass (Bouteloua gra-
cilis) and increasing abundance and productivity of
exotic and native C3 forbs (Alward et al. 1999). These
findings were attributed to climate change and, in
particular, an increase in the spring minimum
temperature.
Long-term ecological studies can detect surprise
events. The only known population of the endangered
Australian heathland shrub, Epacris stuartii, was moni-
tored for several years to quantify vital rates and dif-
ferential fire responses in contrasting microhabitats
(Keith 2002). A sudden episode of 15% mortality of
standing plants was caused by a storm that exposed
the plants to extreme levels of salt spray.The frequency
of equivalent storms was estimated from historical
weather data, allowing evaluation of the combined
impact of fires and storms on the persistence of the
species. A shorter study would have failed to detect the
role that storms play in population turnover. Aware-
ness of this role presented strategic opportunities to
manage more effectively for population persistence
and raised important questions about potential
changes in the frequency of storms under future
climates.
Long-term studies often allow the identification,
quantification and deeper understanding of important
phenomena that are not detectable with a cross-
sectional approach (Wiens 1981; Holmes & Sherry
2001; Kruuk & Hill 2008; Lindenmayer & Cunning-
ham 2011).These include the key ecological processes
that underlie ecological patterns, relationships
between different ecological patterns, links between
sets of interrelated ecological processes and long-term
overall trends that are not reflected by short-term
trends or patterns (Fig. 2). In addition, time is an
independent variable in processes that involve change
such as major natural disturbances, and some ecologi-
cal surprises do not become apparent unless targeted
entities like populations, ecosystems and ecological
processes are studied for a prolonged period (Likens
1989a; Krebs et al. 2001).
doi:10.1111/j.1442-9993.2011.02351.x
 4 D. B. L I N D E N M AY E R ET AL.

1996 1997

2000 2004

B
Grey Shrike-thrush Black-faced Cuckoo-shrike Yellow-rumped Thornbill Common Bronzewing
Estimated probability of detection (%)
Estimated probability of detection (%)

Estimated probability of detection (%)

Estimated probability of detection (%)

25.0 10
16
22.5
14 8 4
20.0
12
17.5 6 3
10
15.0
4 2
12.5 8

10.0 2
6 1
7.5 4 0
1998 1999 2000 2001 2002 2003 2004 2005 1998 1999 2000 2001 2002 2003 2004 2005 1998 1999 2000 2001 2002 2003 2004 2005 1998 1999 2000 2001 2002 2003 2004 2005
Year Year Year Year

Fig. 1. (A) Long-term landscape context changes around a patch of native woodland in the Nanangroe Natural Experi-
ment in southern New South Wales, 1996–2004. (B) Changes in the detection frequency of exemplar temperate woodland
birds inhabiting temperate woodland patches surrounded by maturing stands of Radiata Pine (redrawn and updated from
Lindenmayer et al. 2008).

Understanding complex ecosystem phenomena associated with the El Niño–Southern Oscillation

that occur over a prolonged period (ENSO) interannual climate mode. Long-term eco-
logical studies revealed that two species of desert
Long-term studies are important where there is a rodents, the sandy inland mouse (Pseudomys her-
prolonged lag period until expression of key trends mannsburgensis) and the spinifex hopping-mouse
(Likens 1989a, 1992; Smith et al. 2007; Fig. 2). For (Notomys alexis), were more sedentary and social
example, oceanographic cycles dependent on deep- during wet years when populations increased, but
sea currents occur over decades and perhaps even more mobile and more likely to disperse during the
centuries, making long-term studies critical in marine drier periods (Dickman et al. 2010). Similarly, the
ecosystems (Stommel 1963; Kaiser et al. 2005). Simi- Koonamore Vegetation Reserve in South Australia
larly, arid systems in Australia are distinctively un- was a degraded pastoral lease in 1925 when land was
predictable and long-term studies are required to reserved, fenced and feral animals were removed.
document population irruptions of small mammals This protection allowed a long-term study of vegeta-
following infrequent flooding (Dickman et al. 2010) tion and soil recovery against a background of
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Journal compilation © 2012 Ecological Society of Australia
 VA L U E O F L O N G - T E R M E C O L O G I C A L S T U D I E S
60
50 Precipitation
Hydrogen ion (µeq L–1)
40 trends
30
60
50
40
30
20
10
0 England. This study commenced in 1856 to test the
1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 2010
Year
Fig. 2. Changes in the acidity of precipitation from 1964 to
2009 at Hubbard Brook Experimental Forest, New Hamp-
shire, USA (extended and redrawn from Likens 1989b).
80 years of extreme climatic variability with approxi-
mately decadal cycles (Sinclair 2005).
Long-term studies can be important in quantifying
the effectiveness of conservation management activi-
ties like ecosystem restoration and reconstruc-
tion where prolonged periods might be required
for systems to recover following major human
disturbance. For example, Buckney and Morrison
(1995) implemented a spatially and temporally repli-
cated sampling design to evaluate the success of reveg-
etation treatments on mined Australian coastal sand
plains. They showed that revegetated areas were on a
trajectory towards development of a new ecological
community that differed significantly in species com-
position from pre-mining vegetation and adjacent
un-mined vegetation. Whether the restoration trajec-
tories reach a new stable state or converge over time to
resemble the un-mined community can be resolved
only with continuation of the sampling (Wilkins et al.
2003).
Long-term studies can allow separation of short-
term fluctuations like intra- and interannual variability
from longer-term trajectories (Keeling et al. 1995,
1996; Franklin & Whelan 2009).This understanding is
particularly important where there is substantial tem-
poral variability in ecosystem conditions and popula-
tions of organisms (Wiens 1981; Fuller et al. 1997;
Maron et al. 2005). For example, Lindenmayer and
Cunningham (2011) used a 12-year data set to quan-
tify broad temporal patterns of change in density for a
suite of temperate woodland bird species in south-
eastern Australia and found a significant long-term
trend for increasing abundance for many of the
species, overriding the high levels of year-to-year varia-
tion (Fig. 3; Lindenmayer & Cunningham 2011). This
work was important in that it showed that a number of
species of woodland birds that were thought to be
declining actually exhibited the reverse trend. A long-
© 2012 The Authors
Journal compilation © 2012 Ecological Society of Australia
5
term study of tropical mosquitoes in the Northern
Territory city of Darwin revealed significant seasonal
fluctuations in animal abundance and species diversity
associated with onset and cessation of the wet season.
However, there was striking predictability in the
annual cycles of mosquito assemblages, irrespective of
the irruptive dynamics of some individual species
(Franklin & Whelan 2009).
Perhaps, one of the most famous and enduring long-
term ecological studies is the Park Grass Experiment
at Rothamsted Experimental Station in Hertfordshire,
effect of fertilizers on hay yields, but has since been
modified to include other treatments. Results over this
long period have demonstrated that: (1) conventional
field trials probably underestimate threats to plant
biodiversity from long-term changes, such as soil acidi-
fication; (2) plant species richness, biomass and soil
pH are strongly interrelated; (3) competition between
plants can magnify the effects of climatic variation on
communities; and (4) local evolutionary changes can
occur under different selection pressures (Silvertown
et al. 2006).
Providing core ecological data for developing
theoretical models and to parameterize and
validate simulation models
Empirical field data are fundamental for testing eco-
logical theory (Shrader-Frechette & McCoy 1993;
Franklin & MacMahon 2000). In Australia, a wide
range of evolutionary and ecological theories have
been tested through classic long-term behavioural
studies such as those on the superb fairy-wren
(Malarus cyaneus) (Cockburn et al. 2008) and the
sleepy lizard (Tiliqua rugosa) (Bull & Burzacott 2002,
2006; Leu et al. 2010). A long-term and ‘large-scale’
study conducted in the tropical moist forests of
Panama since 1923 has provided data to test models
in community ecology about the maintenance of
diversity. This study led to the establishment in
1980 of the original 50-ha plot for long-term moni-
toring of its trees (Condit 1995). The programme
of research has expanded to plant physiology, canopy
biology and animal ecology, and there is now a
network of over 40 large permanent plots world-
wide (http://www.ctfs.si.edu/group/About/). The
complete enumeration and spatial mapping of all
trees stimulated the development of the unified
neutral theory of biodiversity and biogeography
that seeks to explain the relative abundance of
species in ecological communities (Hubbell
2001, 2006). The fully archived data are provided
as a shared resource for all ecologists (http://
knb.ecoinformatics.org/knb/metacat/nceas.298/
nceas).
doi:10.1111/j.1442-9993.2011.02351.x
 6 D. B. L I N D E N M AY E R ET AL.

Brown Treecreeper Diamond Firetail Restless Flycatcher

70 20.0 40
17.5 35
60
15.0 30
Reporting rate (%)

Reporting rate (%)

Reporting rate (%)

50 12.5 25

40 10.0 20
7.5 15
30
5.0 10
20 2.5 5
0.0 0
10
1998 2000 2002 2004 2006 2008 1998 2000 2002 2004 2006 2008 1998 2000 2002 2004 2006 2008
Survey year Survey year Survey year

Grey Shrike-thrush Crested Shrike-tit

80 40

70 35

30
60
Reporting rate (%)

Reporting rate (%)

25
50
20
40
15
30 10

20 5

10 0
1998 2000 2002 2004 2006 2008 1998 2000 2002 2004 2006 2008
Survey year Survey year

Fig. 3. Temporal changes in the reporting rate of a range of temperate woodland birds in south-eastern Australia (redrawn
from Lindenmayer & Cunningham 2011).

Long-term studies are critical for generating the recurring at subdecadal to multidecadal intervals.
kinds of data needed for simulation modelling – an They constructed a stochastic population model
increasingly important part of ecology and environ- parameterized by data from a 20-year annual census of
mental management (Burgman et al. 1993; Canham survival, growth and reproduction (Keith et al. 2007).
et al. 2003; Pople et al. 2010). As an example, long- The count sampled demographic responses to three
term ecological data were used to build spatially successive fires, a number barely adequate to charac-
explicit models of kangaroo populations (Pople et al. terize the variability of fire-driven responses in vital
2007), small desert mammals (Dickman et al. 2011) rates. Combined results from the model and surveys
and arboreal marsupials (Lindenmayer & McCarthy produced evidence of a decline in the population,
2006). Lindenmayer and Possingham (1995) and Lin- alerting managers of the need to diagnose the cause
denmayer and McCarthy (2006) coupled long-term and explore alternative management responses to con-
empirical data and simulation modelling to assess the serve this functionally important species (Regan et al.
viability of populations of the nationally endangered 2011).
Leadbeater’s possum (Gymnobelideus leadbeateri) and Long-term studies are also critical for informing
guide the expansion of a reserve system for the species ecological genetic models aimed at understanding the
(Department of Sustainability and Environment ability of species to evolve and/or adapt to environ-
2008). The ongoing collection of long-term field data mental changes (Hoffmann & Sgro 2011). Much of
has also facilitated a cycle of testing and continuous our understanding of evolutionary potential comes
upgrading of simulation models (Lindenmayer & from long-term studies like those involving the Gal-
McCarthy 2006). apagos finches (Grant & Grant 2002) and Soay sheep
Long-term studies are particularly important for (Ovis aries) (Ozgul et al. 2009). These data sets have
parameterizing models of species with slow population been essential for testing models of adaptation and for
turnover or where turnover is governed by infrequent developing an understanding of how trait interactions
events. Regan et al. (2011) reported that survival, and patterns of natural selection operate and deter-
growth and reproduction of the long-lived Australian mine the long-term viability of populations and
grass tree, Xanthorrhoea resinosa, are influenced by fires species.
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 VA L U E O F L O N G - T E R M E C O L O G I C A L S T U D I E S
Acting as platforms for collaborative studies,
thus promoting multidisciplinary research
Long-term studies can often be the stimulus for other
investigations either at the same site or at other sites.
For example, a network of long-term monitoring sites
maintained in the wet ash eucalypt forests of south-
eastern Australia has formed the framework for a series
of other ecological studies (Lindenmayer 2009a).
Similar catalytic effects have occurred around long-
term studies of vertebrate population dynamics in arid
Australia (Letnic & Dickman 2010).There are numer-
ous examples of this from around the world such as the
long-term studies of mammal population cycles in the
boreal forests of Canada (Pastor et al. 1993; Krebs
et al. 2001; Persson et al. 2009). An important aspect
of co-located investigations is the instigation of mecha-
nistic studies that account for the long-term patterns
that have been identified (e.g. Pastor et al. 1993; Krebs
et al. 2001; Persson et al. 2009).
Long-term ecological studies at a given site can
become particularly powerful when a multidisci-
plinary approach is taken. The value of long-term
ecological research can be significantly increased
when it is connected to physicochemical information
to help interpret ecological phenomena (Bormann &
Likens 1979; Malhi et al. 2008; Hutley et al. 2011).
For example, long-term studies on a lake system in
Washington, USA highlighted the impacts on aquatic
ecosystems of increasing levels of pollutants
(Edmondson 1991). In other cases, linking long-term
ecological studies with work in other disciplines such
as medicine can yield profound insights. As an
example, based on work in south-eastern Australia,
Johnston et al. (2011) uncovered important interrela-
tionships between patterns of vegetation cover,
altered burning regimes and changes in human
health outcomes. Indeed, we suggest that co-located
multidisciplinary studies that connect long-term eco-
logical research with other bodies of work and allow
storage and sharing of the data, under appropriate
conditions by ecologists and others, will become
increasingly critical in the future for ecology and
the knowledge it provides for natural resource man-
agement. These projects work best if the communi-
ties openly cooperate and share knowledge and data
to build on collaboratively. The facilities supported
under Australia’s Terrestrial Ecosystem Network aim
to establish the resources for this type of science.
Long-term studies at one location can stimulate
others to carry out similar investigations at other
locations. The pioneering work that established the
first 50-ha permanent forest plot (Hubbell & Foster
1983, 1986) at Barro Colorado Island and enumer-
ated all trees with a diameter at breast height of at
least 1 cm has grown into a global network of forest
research plots united by a common methodology and
© 2012 The Authors
Journal compilation © 2012 Ecological Society of Australia
7
committed to the study of function and diversity in
tropical and temperate forests. The multi-institutional
network monitors the growth and survival of about
3.5 million trees of approximately 7500 species
(http://www.ctfs.si.edu/group/About/). Australia rec-
ently established a 25-ha plot in far North Queen-
sland, and will soon be joining the above-mentioned
network.
In this way, long-term monitoring plots become a
valuable component of a much larger network and
substantial statistical power is provided to look for
common trends. The International Tundra Experi-
ment (ITEX) provides one example where monitoring
and experimental plots have been located in tundra
and alpine areas throughout the world and data have
been used in meta-analyses to investigate vegetation
and soil responses to warming and changing precipi-
tation patterns (Arft et al. 1999). An Australian com-
ponent of this network has been established (Jarrad
et al. 2008).
For a long-term study to stimulate parallel investi-
gations in other locations, data, meta-data and
methods must be made available in a form that can be
discovered, accessed, used with confidence and cited
as a source. Indeed, proper attribution of long-term
data remains a difficult and unresolved challenge (see
Likens & Lindenmayer 2011). Such a rich data
resource is the exception in ecology, and in most areas
of science, as shown in recent international reviews
(Tenopir et al. 2011). A number of national research
agencies (e.g. the US National Science Foundation)
are establishing projects to allow data to be stored and
used under suitable licensing conditions (e.g. the
DataNET and DataONE projects in the USA and
TERN in Australia). The newly developing ANDS
(Australian National Data Service) may become
another opportunity for storage and wider access to
long-term ecological data in Australia.
Providing data and understanding to support
evidence-based policy, decision making and
management of ecosystems
Long-term observations are critical for making
informed decisions in many environmental manage-
ment contexts. For example, long-term population
studies of large macropods in Australia underpin the
kangaroo harvesting industry by setting ecologically
sustainable annual quotas. Population data are gath-
ered regularly, both through aerial survey and harvest
returns, and indicate marked fluctuations in density
in response to droughts and wet years. Long-term
data from various sources are integrated to guide
harvest quotas based on population trends, harvest
levels and climatic conditions (Pople et al. 2007,
2010).
doi:10.1111/j.1442-9993.2011.02351.x
8 D. B. L I N D E N M AY E R ET AL.
Studies into the effects of fire regimes on vegetation
and fauna in fire-prone Australian savanna systems
have been ongoing since the early 1970s, and have
included replicated experimental designs (Andersen
et al. 2003; Russell-Smith et al. 2003), extensive
assessments of ambient fire regime effects (Prior et al.
2009; Russell-Smith et al. 2010; Woinarski et al.
2010), spatial application of plot-based models to
predict landscape-scale change (Russell-Smith et al.
unpubl. data 2011) and a broadened research focus
addressing carbon sequestration and related issues
(Liedloff & Cook 2007; Murphy et al. 2010).Together,
these bodies of long-term work have created novel
opportunities to develop payments to regional land
managers for ecosystem service arrangements based
on enhanced fire management of biodiversity and
carbon resources (Whitehead et al. 2008). In addition,
faunal sampling since 1996 in one reserve has demon-
strated a precipitous decline in small mammal popu-
lations, with both plot-level species richness and total
abundance decreasing by greater than 50% in about
10 years (Woinarski et al. 2010). While the assembled
data implicated a contributory role of frequent fire in
the observed collapse, it is likely that other factors also
were involved such as predation by cats (Felis catus),
other feral animals and disease. These observations
highlight the need for ongoing targeted studies to help
disentangle potential multiple causal agents and create
the basis for informed and effective biodiversity con-
servation in northern Australia.
Long-term studies can be pivotal for informing
evidence-based environmental legislation such as
laws to control air and water pollutants. For instance,
legislation regulating the release of pollutants like
carbon dioxide or sulphur dioxide is typically imple-
mented without a clear view of the time frame of the
response. In the USA, the Clean Air Act Amend-
ments of 1970 and 1990 mandated a reduction in
particulate and sulphur dioxide emissions from
industrial combustion of coal and oil to reduce the
amount of air pollution and acid rain falling on
eastern North America. Emissions of sulphur dioxide
to the atmosphere are a major precursor in the for-
mation of acid rain. No one had expected that pro-
longed data collection would be needed to test the
effectiveness of this legislation, but it required
18 years of data on precipitation chemistry in the
Hubbard Brook Experimental Forest in New Hamp-
shire, USA to show that the acidity of precipitation
had actually decreased (Fig. 2) (Likens 1989b, 1992)
in response to these mandated reductions in emis-
sions. This ostensibly ‘simple’ result was needed by
policy makers dealing with the problem of acid rain,
but took a long time to obtain (Likens 1992).
Finally, in the present context of rapid climate
change, one of the most important examples of a long-
term study is the work initiated in 1958 on Mauna
doi:10.1111/j.1442-9993.2011.02351.x
Journal compilation © 2012 Ecological Society of Australia
390
380
CO2 concentration (ppm)
370
360
350
340
330
320
310
1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 2010
Year
Fig. 4. The Mauna Loa (Hawaii, USA) record of atmo-
spheric CO2 concentrations (redrawn from http://www.
scrippsco2.ucsd.edu).
Loa, Hawaii that has demonstrated marked increases
in the concentration of atmospheric carbon dioxide
(Keeling et al. 1995, 1996) – a trend that continues
today and is now investigated through a global network
of stations (Battin et al. 2009) (Fig. 4).
DISCUSSION
Despite the many values of long-term ecological
studies (summarized in Table 1), they are not particu-
larly common in Australia (reviewed by Youngentob
et al. unpubl. data 2011) or elsewhere around the
world. Reasons include lack of funding, the relatively
short working lifespan of the researchers who cham-
pion the study and a failure of scientists to communi-
cate effectively the array of values of long-term studies,
including their key practical value in informing man-
agement and decision making. An additional reason is
the social and political fascination with new and inno-
vative projects – a problem that has sometimes led to
long-term studies and especially long-term ecological
monitoring being dubbed ‘Cinderella science’ (Nisbet
2007). These problems can lead to a lack of ‘official
recognition’ in government and research funding
bodies of the unique and essential nature of long-term
studies (Lovett et al. 2007; Lindenmayer & Likens
2010), in turn leading to difficulties in obtaining the
necessary funding to maintain them (Lindenmayer &
Gibbons 2012). These problems, coupled with the
considerable resources needed to maintain long-term
studies, highlight why it is critically important for such
work to be well motivated, efficiently designed and
implemented, relevant to management and policy
making (Russell-Smith et al. 2003) and where possible
demonstrated to be cost-effective. For example, the
estimated annual cost of implementing legislative
reforms on acid rain in the USA was $8-9 billion
per year but these reforms delivered positive annual
© 2012 The Authors
 VA L U E O F L O N G - T E R M E C O L O G I C A L S T U D I E S
Table 1. Summary of some of the key values of long-term ecological studies
Value
Quantifying ecological responses to environmental change
Understanding complex ecosystem phenomena that occur over a
prolonged period
Providing core ecological data for use in developing theoretical
models and in parameterizing and validating simulation models
Acting as platforms for collaborative studies, thus promoting
multidisciplinary research
Providing data and understanding to support evidence-based
policy, decision making and management of ecosystems
environmental benefits valued at $101–119 billion
(Oreskes & Conway 2010). We are not aware of any
careful analysis of the total research costs for the
underlying, long-term studies leading to the identifi-
cation of the acid rain problem and its causal factors.
However, based on the cost of the Hubbard Brook
Ecosystem Study that first identified the problem of
acid rain in North America, and the cost of a major US
programme called the National Acid Precipitation
Assessment Program, we estimate a cost of less than
$100 million per year in the USA, representing an
extraordinary level of cost-effectiveness relative to the
scale of the problem.
As valuable as long-term ecological studies clearly
are, it is also important to recognize that some may
reach a stage when some key questions have been
answered and it is appropriate to reduce the level of
investment in them. However, this will often not be
the case because of the relative rarity of long-term
studies and the many values generated from main-
taining such investigations. Moreover, careful exami-
nation should be undertaken of the completeness of
the knowledge obtained from existing long-term
studies as valuable and unexpected insights can often
be generated from maintaining data collection. For
example, the Mauna Loa survey of levels of atmo-
spheric carbon dioxide (Keeling et al. 1995, 1996)
was threatened with termination six times (Linden-
mayer & Likens 2010) with some reviewers of the
project insisting that enough was known about trends
and patterns. Others successfully argued that it was
important to continue gathering additional informa-
tion which now supports the recognition of rapid and
continuing increases in atmospheric carbon dioxide.
There are many environmental issues which have
generated mandatory long-term monitoring as
required under formal plans of management, or for
ongoing monitoring of carbon stocks in natural
systems to acquire data used for assessing climate
change effects (Willis & MacDonald 2011).
© 2012 The Authors
Journal compilation © 2012 Ecological Society of Australia
9
Examples
Likens et al. 1970; Schindler et al. 1985; Keeling
et al. 1995, 1996; Chave et al. 2003; Metcalfe
and Bradford 2008; Willis et al. 2008; Dickman
et al. 2010; Russell-Smith et al. 2010
Keeling et al. 1995, 1996; Lindenmayer and
Cunningham 2011
Burgman et al. 1993; Lindenmayer and McCarthy
2006; Regan et al. 2011
Bormann and Likens 1979; Krebs et al. 2001;
Hubbell 2002; Lindenmayer 2009a,b
Likens 1992; Russell-Smith et al. 2003; Sutherland
et al. 2004; Woinarski et al. 2010
If the aim of an ecological study is to quantify a
particular long-term process, as the study proceeds
there may be diminishing returns, for example, in
improving the precision of an estimate. Under these
circumstances, it may be more fruitful to direct
resources into other studies. Decision analysis can help
determine optimal trade-offs for cost-efficient long-
term ecological research (White 2005; Chades et al.
2008). However, such decisions become complex
when detection of surprises and other less tangible
goals are an important component of the research.
Such decisions are also complex for surveillance (non-
question-driven) long-term studies which can some-
times lead to the discoveries of important new
phenomena or lead to valuable discoveries that were
not part of the original objectives of the investigation
when first instigated (Franklin & Whelan 2009; Wintle
et al. 2010). The discovery of eggshell thinning result-
ing from pesticide use (Grier 1982) is one of many
examples.
The future of long-term ecological studies
Recent trends in ecological research may make it dif-
ficult to maintain long-term studies, with much
emphasis now placed on approaches using meta-
analysis, data mining and modelling. These can be
powerful and are generally cheaper and quicker to
generate trends and publications than traditional
field-based, empirical ecology. This makes it difficult
for field ecologists running long-term studies to
compete with modellers and data summarizers when
funding success is often based on publication metrics
(Lindenmayer & Likens 2011). In addition, in some
areas of ecological research, it appears to be increas-
ingly difficult to publish empirical, place-based
studies compared with modelling, meta-analysis and
data mining. Bormann et al. (2007) and Westgate
et al. (unpubl. data 2011) showed that empirical
doi:10.1111/j.1442-9993.2011.02351.x
10 D. B. L I N D E N M AY E R ET AL.
examples of adaptive management are significantly
harder to publish in the international peer-reviewed
literature and are also significantly less cited than
non-empirical opinion pieces and essays on the same
topic. Yet, empirical data are the raw materials essen-
tial for intelligent parameterization of models and
meta-analysis. Without ongoing support for long-
term ecological studies, meta-analyses and systematic
reviews will have fewer cases to review and analyse,
important ecological discoveries may be overlooked
and evidence-based environmental policy and man-
agement (e.g. Sutherland et al. 2004; Collaboration
for Environmental Evidence 2011) will become
harder to achieve (Lindenmayer & Likens 2011).
Moreover, we suggest that without long-term ecologi-
cal studies, it will not be possible to tackle effectively
key issues like the management of ecosystems for
their key services and quantify and mitigate the
effects of environmental change on ecosystems and
on biodiversity, including climate change.
To ensure the maintenance of long-term ecological
studies, we believe it is important to maintain pub-
lishing outlets for empirical field-based ecology,
develop new systems of recognition that reward
empirical ecologists for the use of their data by
others (e.g. Likens & Lindenmayer 2011; Reichman
et al. 2011) and re-craft funding schemes to empha-
size collaborative partnerships among field-based
ecologists, theoreticians and modellers as well as
between ecologists and workers in other fields as part
of multidisciplinary research. Moreover, funding
schemes in ecology per se need to better acknowl-
edge the importance and manifold values of long-
term studies and ensure that this translates to
financial support for such work. Trends in this direc-
tion are now included in the US National Science
Foundation and under the Terrestrial Ecosystem
Research Network in Australia (albeit only until 2014
for the latter at this stage).
ACKNOWLEDGEMENTS
This paper arose as a result of a workshop sponsored
by the Terrestrial Ecosystem Research Network
(TERN). We thank Professor M. McGeoch and Pro-
fessor S. Ormerod for insightful comments that con-
siderably improved an earlier version of this paper.
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