Nordic Society Oikos

Oikos, 50 Years of Ecology

Author(s): Jan Lindström, Per Lundberg, Esa Ranta and Veijo Kaitala
Source: Oikos, Vol. 87, No. 3 (Dec., 1999), pp. 462-475
Published by: Wiley on behalf of Nordic Society Oikos
Stable URL: https://www.jstor.org/stable/3546810
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OIKOS 87: 462-475. Copenhagen 1999

Oikos, 50 years of ecology

Jan Lindstrom, Per Lundberg, Esa Ranta and Veijo Kaitala

F < . t Lindstrom, J., Lundberg, P., Ranta, E. and Kaitala, V. 1999. Oikos, 50 years of
4" %ecology. - Oikos 87: 462-475.

J. Lindstrom, Dept of Zoology, Univ. of Cambridge, Downing Street, Cam

CB2 3EJ (j.lindstrom@zoo.cam.ac.uk). - P. Lundberg, Dept of Theoretical Ecology,
Lund Univ., Ecology Building, SE-223 62 Lund, Sweden. - E. Ranta and V. Kaitala,
S . Ad Integrative Ecology Unit, Div. of Population Biology, Dept of Ecology and Systematics,
P.O. Box 17, FIN-00014 Univ. of Helsinki, Finland (present address of VK: Dept of
Biological and Environmental Science, Univ. of Jyviskylk, P.O. Box 35, FIN-40351
Jyvdskyld, Finland.)

Scandinavian (or more correctly: Fennoscandian) ecol-
ogy has come to age with the Nordic Ecological Society
Oikos reaching its 50-year anniversary this year
(Maimer and Ranta 1999). From the very beginning,
the founding fathers decided to launch an ecological
journal, Oikos. The anniversary year of the Society has
been celebrated by inviting a number of specialists to
write overviews on the important developments in their
own research fields of ecology. This resulted in 20
articles published as Minireviews in Oikos and Ecogra-
phy during 1999. The scope of this essay is to put the
journals and the reviews into the context of the devel-
opment of ecology as a discipline.
The launching of the journal Oikos coincides with
the maturation of ecology to a solid scientific discipline.
The topics discussed on the pages of this journal,
already from the first volume onwards, had a tight
connection to the development of ecology in Britain
and the USA. In the early years, the term ecology was
just a name for a scientific discipline. In the years to
come, the land-mark decade being the 1960s, ecology
has been variously associated with the increasing envi-
ronmental concern in political awareness. Firstly with
the pollution threat and lately with the loss of habitats
- especially in temperate areas and in the tropics - and
the associated biological diversity. These days, the term
ecology appears in the names of various commercial
products from toilet paper to computer displays.
Surely, every educated person has some mental icon for
the term ecology.
For the ecologists, the term ecology still adheres to
the classical definition of ecology: the interplay of indi-
viduals and their environment. This is a rather broad,
almost meaningless, definition but as Ernst Mayr (1963)
remarked "There is no area of biology in which evolu-
tion has not served as an ordering principle". In ecol-
ogy, this is probably now more true than ever as
ecology incorporates sub-fields like evolutionary and
behavioral ecology. It also appears that contemporary
population (quantitative) genetics has more to do with
ecology than with genetics.
Oikos 1949-1999
Oikos is a multidisciplinary ecological journal. Even a
casual comparison of its contents with those of Ecology
or Journal of Animal Ecology and Journal of Ecology
proves this. However, we were curious to see what has
happened to Oikos during its first 50 years (87 vol-
umes). We decided to do what ecologists often do: to
collect data. We selected three sampling periods, the
first 10 years (1949-1958), the middle years (1969-
1978) and the last decade (1989-1998). For each pe-
riod, a total of 50 articles were drawn at random from
the articles published in each period. Of course, the
sampling coverage of the whole content of the journal
was higher in the early years when articles were fewer
and longer than during the past decade or two, when

This is an invited Minireview on the occasion of the 50th

anniversary of the Nordic Ecological Society Oikos.

Copyright ?) OIKOS 1999

ISSN 0030-1299
Printed in Ireland - all rights reserved

462 OIKOS 87:3 (1999)

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the number of volumes and printed pages per year in
Oikos have kept increasing. We also did a parallel
sampling in Ecology, a journal similar in scope to
Oikos. The articles thus collected were scored into eight
groups (originally we had more categories, but the
picture emerging is more clear with the current group-
ing) based on their content.
The categorizing is certainly a biased and subjective
procedure and so is the classifying of the individual
papers. One can nevertheless draw a few conclusions
(Fig. 1). First, there are temporal changes in the journal
profiles over the decades. For example, the early years
were more dominated by articles on ecosystem and
community ecology than later on. Also, pure theory
was scanty in the first volumes, as was also behavioral
ecology (officially born in late 1970s), but both are
present in the 1970s and 1990s. The two journals differ
surprisingly little in their profiles (Fig. 1), with perhaps
the only exception being that Oikos tends to publish
less ecosystem research than Ecology. Perhaps the
match in profiles between the two journals is under-
standable. They are "general" ecology journals, and
can both be taken as representative samples of what
high-quality manuscripts there are available in the en-
tire domain of ecology.
We were also curious to see what has happened to
the authorship composition of Oikos during all these
years. The launching of the journal was entirely a
Scandinavian enterprise. However, the ethnical origin
of writers soon started to diverge. In the first five
volumes, Scandinavian authors contributed 46% of all
published articles, while in volumes 6 to 10 the corre-
sponding figure was as low as 12%. Since then, despite
the fact that due to its proximity Oikos is a natural
choice for many Scandinavians, the proportion of non-
Scandinavian authors has remained high. Perhaps the
biggest change, in terms of Oikos - and Scandinavian
ecology itself - becoming international began in the
late 1970s when several esteemed North American ecol-
ogists started to contribute to this journal. This was due
to active journal policy, the agents being Per Brinck
and Staffan Ulfstrand.
To complete the exercise of data collection we also
scanned the list of references, and scored the occurrence
of citations to Scandinavian ecologists, in ecological
textbooks of the 50-year period. We did this well aware
that the figures do not exactly tell the contribution of
Scandinavians to the world's ecological scientific litera-
ture. To us, the emerging figures were rather sobering:
Clarke (1954) 3.6%, Odum (1959) 2.1%, Krebs (1972)
1.3%, Pianka (1978) 0.8% and Begon et al. (1996) 2.9%.
We cannot avoid speculating: either Scandinavians do
poor ecology (which we hope is not true) or textbook
writing is strongly biased. Actually, a recent Finnish

Oikos

1949- 1958 1969- 1978 1989- 1998

Theory

Autecology \ \ -

Physiol ecol

Behavioral ecology

Life-history

Plant-animal interaction

Community

Ecosystems

Ecology

Theory

Autecology

Physiol ecol

Behavioral ecology
Life-history

Plant-animal interaction
Fig. 1. Publication profile C t_ M
(eight categories) of Gikos and Community
Ecology in three different Ecosystems - - ,
10-year periods. The data are
random samples of articles 0 10 20 30 40 0 10 20 30 0 10 20 30 40
published in the two journals
during each of the three Frequency, %
1 0-year periods.

OIKOS 87:3 (1999) 463

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ecology textbook (Hanski et al. 1998) shows indeed a
similarly strong bias in the selection of examples: 35%
of the references were by Scandinavian ecologists.
To include the second of the Nordic Society Oikos'
journals, Ecography, in this initial comparison of the
development of the contents between the multidisci-
plinary journals Oikos and Ecology would not be fair;
first because Ecography has only existed since 1978
(and thus cannot be included in the first two time
periods in Fig. 1), and second because Ecography has a
different profile, centering on patterns in ecology more
than processes (spatial and temporal patterns, natural
history of species). However, in the second part of our
review all seven papers published in Ecography are
included.
The anniversary essays
The essays that have appeared in Oikos and Ecography
during this anniversary year have covered a wide range
of topics, emphasizing some, omitting others. We will
here rhapsodically summarize them and attempt to put
them into the historical and prospective context of this
paper.
Are generalizations possible?
Most ecologists would probably subscribe to the idea
that ecology is rich in theory, but poor in laws. It all, of
course, depends on how "law" is defined and attempts
have also recently been made to find adequate defini-
tions of the concept that are appropriate for ecology
(e.g. Pickett et al. 1994). Lawton (1999) asks to what
degree generalizations are possible in ecology and
whether such generalizations should be called "laws".
Lawton concludes that the laws in ecology are (and
must be) different from the deep universal laws in, for
example, physics. Having said so, Lawton nevertheless
identifies a set of fundamentals that underpin all eco-
logical systems; for example, the first and second laws(!)
of thermodynamics, Darwinian natural selection, the
physical principles governing diffusion and transporta-
tion of matter and some fundamental mechanics (re-
garding, e.g., flight, bone structures, etc.). Hence, some
things are inevitably in common to all organisms, but is
it possible to go beyond this?
Lawton seeks an answer by taking a closer look at
three areas of ecological inquiry for which a substantial
body of theory exists. The areas are population dynam-
ics, community ecology and macroecology. First, Law-
ton admits that all organisms, even all local
populations, are unique and therefore at one level
escape generalization. This is rather trivial. On the
other hand, there is a limited number of qualitative
dynamic properties. The exact nature of the dynamics
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of a particular species or population will be, however,
contingent on a suite of special circumstances, often
unique in space and time.
Lawton argues that communities are more problem-
atic. They are harder to define, it is arguable what
patterns are to be explained and understood by general
theories. Communities are so complicated that contin-
gent theory does not seem to work very well, but they
are also small enough such that global patterns would
not appear. That is why macroecology has better
chances to reveal order and generality. It attempts to
embrace major, statistical patterns of distributions,
abundances, species richness and alike. It also empha-
sizes the interplay between local and global scales, a
common theme throughout turn-of-the-millennium
ecology (cf. for example, Petersen et al. (1999) and
Huston (1999) below). Macroecology is full of consis-
tent patterns across scales and taxa. Perhaps it is the
scale-dependence that is both the beauty and the curse
of ecology. Lawton is a modern ecologist, acknowledg-
ing both the complexity and uniqueness of ecological
systems, as well as the scientific endeavor of finding
general laws and their mechanisms. It remains to be
seen how bold and daring the ecologists of the next
millennium are going to be in generalizing the living
world.
In their respective reviews, Persson (1999) and Polis
(1999) deal with what Lawton thinks is the most
difficult scale - general patterns and processes in com-
munities. Polis appears to prove Lawton's case. Perhaps
ecologists have become so engaged with sophisticated
measuring devices, complicated mathematical models
and a wish to find biotic explanations for everything
that they can not see the wood for the trees. We believe
that Polis would subscribe to this statement, although
he does not explicitly say it in the essay on trophic
cascades and the factors controlling primary productiv-
ity. The "Green world, or the HSS" hypothesis sug-
gested by Hairston et al. (1960) 40 years ago sparked a
never-ending quest for the explanation of the green
world. The world is green, they argued, because herbi-
vores are kept under control by their predators. Should
predators be absent, the herbivores would over-exploit
the plants and the world would turn yellow or red.
Fretwell (1977), and later Oksanen et al. (1981), sug-
gested that this is exactly what we can see if we follow
gradients of primary productivity. Along such a gradi-
ent, the primary productivity would be able to sustain
food chains of different length such that the world
would be green in systems with odd-numbered links.
Polis takes us back to square one and asks whether
the complex interactions in natural communities really
have the power of producing the observed patterns of
"greenness" around the world. No, says Polis and seeks
more parsimonious explanations. The list of such expla-
nations contains sunlight and climate together with
catastrophes and large-scale abiotic disturbances, and
87:3 (1999)
overall nutrient and water availability. Reviewing ob-
servations from both aquatic and terrestrial ecosystems,
Polis concludes that herbivores in aquatic systems actu-
ally may have the potential of influencing plant biomass
and functioning significantly, whereas terrestrial herbi-
vores rarely can. Polis also lists the reasons why we
indeed would not expect herbivores to have the capac-
ity of regulating plant biomass. These include plant
defenses and self-regulation. According to Polis, PPPP
(predators, parasitoids, parasites, and pathogens) do
not seem able to do the job. With a few exceptions,
however. Under some rare circumstances the conditions
are "right" for community-level cascades, for example
in some aquatic systems with rather simple food webs.
There are a number of reasons why aquatic systems are
expected to be different from terrestrial ones in this
respect. Size-ratios across trophic levels are very differ-
ent in aquatic and terrestrial communities and this has
consequences for, e.g., consumption rates and life-cycle
relationships.
In Polis' mind the world's ecosystems are too hetero-
geneous spatially and temporally to allow for any gen-
eralizations about the impact of herbivores on plant
biomass.
Persson (1999) is more optimistic, given that we
include the relevant biological features of the systems
under study. Persson, like Polis, is interested in trophic
cascades. Together with the "bottom up-top down"
dichotomy, it is an attempt to put a label on a seem-
ingly widespread phenomenon, particularly in aquatic
ecosystems. The often used definition of a trophic cas-
cade is the propagation of indirect mutualism between
non-adjacent levels in a food chain. Although there
seems to be ample support for trophic cascades (Polis
would not agree), they appear to be confined to aquatic
systems. The reasons for this may be two-fold. First,
aquatic food chains are often simpler and are defined
for a common spatial and temporal scale. Second,
attempts to find trophic cascades in terrestrial systems
may be flawed because too much focus has been on
plant-insect food chains, omitting mammalian herbi-
vores. But Persson argues that the problems with find-
ing trophic cascades may go beyond the measurement
problem. The basic idea in trophic cascade theory is
that the effects occur across trophic levels. In the
"exploitation ecosystems" (Fretwell 1977, Oksanen et
al. 1981) all species and populations at each trophic
level can be lumped together. Persson argues that this is
not necessarily the case and lists a number of reasons
for this. One particularly important problem is that
productivity (a corner stone in trophic cascade theory)
is not well defined. Lumping species together and creat-
ing trophic levels may also be problematic because the
pathways of energy and matter through the system are
easily modified by direct or indirect species interaction
within levels. This is why Polis thinks that community-
wide cascades are rare and species cascades are more
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common. Trophic level omnivory (vertical heterogene-
ity) will also make the identification of trophic levels,
and therefore trophic cascades, impossible. Even more
disturbing is life-history omnivory, i.e., that organisms
may pass and use several trophic levels throughout
their lifetime. They grow and use different habitats
within the system at different life-stages.
Persson also stresses the fact that there is more to
trophic dynamics than cascades. Adaptive foraging,
nutrient recycling, size-structured dynamics and habitat
selection problems all add to indirect effects in both
aquatic and terrestrial systems.
Macroecology
Leaving populations and communities behind, Lawton
(1999) puts more hope to what has been called macroe-
cology. Several of the anniversary reviews address
problems within that framework. Compared with, say,
many deterministic physical systems, ecological com-
munities and ecosystems are immensely complex.
Macroecology takes up this challenge and asks: "Are
there actually general, simple rules for creating this
complexity?". In macroecology, these rules are most
often sought for by documenting broad statistical pat-
terns in communities and ecosystems.
"Classic" macroecology
Brown (1999), a co-inventor of the term "macroecol-
ogy", is one of the anniversary reviewers of this topic.
He stresses that macroecology is rather a research
program and a way of thinking than a subdiscipline of
ecology. According to Brown, the essence of macroecol-
ogy is two-sided. There is the empirical part, the actual
documentation of patterns. The second part is more
theoretical and its task is to formulate hypotheses. On
the whole, macroecology has been more successful in
the first part. Many large-scale patterns have been
found but finding the mechanisms underlying these
patterns has proven to be more difficult. Perhaps this is
not very surprising as there is a great gap between
understanding patterns and understanding processes in
general. A point of comparison could be, for instance,
population cycles. There the pattern is clear and can be
adequately described but finding the actual causative
mechanisms has been a hard challenge in population
ecology [Stenseth (1999) provides an up-to-date view on
this topic in his anniversary review]. An obvious reason
for choosing this evidently difficult approach in
macroecology is its large scale: most systems cannot be
manipulated at the ecosystem level. Yet this is the scale
of climatic change, for instance. The positive message
from macroecology is that finding organized patterns
gives promises of the general mechanisms to exist as
(1999) 465
well. In looking forward to the future of macroecologi-
cal research, Brown emphasizes the need for models to
help in developing the theoretical background.
One of the first large-scale patterns to be discovered
by early ecologists was that the number of species
increases with decreasing latitude. That is, the tropics
are much more species rich than areas close to the
poles. The list of possible explanations for this seem-
ingly general pattern is long. The anniversary review within
by
Rohde (1999) is a thorough and critical examination of
the hypotheses suggested so far.
Before us, we have a good "macroecological" pattern
- the number of species within a given taxon increases
as we go from the poles towards the tropics. Or does it
really? For a number of taxa (e.g., birds, non-flying
mammals, marine fish) it does. But there are numerous
examples where this pattern is absent or unclear, e.g.,
Old World sawflies and galling insects peak at interme-
diate latitudes.
According to Rohde, many of the suggested explana-
tions either lack logical consistence or are not complete
enough or are simply refuted by data. A common
problem to many of them is also that they can discover
correlations between, e.g., certain environmental vari-
ables and species richness, but they do not "explain"
the patterns. Interesting patterns can emerge when par-predicting species diversity? Can we derive any reason-
asites are studied because they cannot be disentangled
from their hosts. If (potential) host species richness
increases towards the equator (for whatever reason), so
should parasite species richness. And it generally does,
but host species richness and parasite diversity change
at different rates, indicating, among other things, that
the patterns we see are not at equilibrium. Rohde
concludes that assuming "saturated" communities is
flawed. Places on earth where energy input is high
(presumably the tropics in most cases), the speed of
evolution is high (promoting speciation). Those places
have also been there unaffected by major disruptions
(e.g., glaciation) for a long time. Consequently, the
pole-near areas should not be saturated and therefore
species poor relative to the tropics. Hence, "effective
evolutionary time" is, according to Rohde, the most
likely explanation for the observed gradients.
Despite the title of the review by Gaston and Black-
burn (1999), they do not actually criticize macroecology
but rather review the criticism presented. By selecting
the topics which are, in their view, most commonly (1999) "laws") prevail in ecology has been frequently
misunderstood they summarize the defense of macroe-asked in the anniversary reviews. Rosenzweig and Ziv
cological approach against its criticism. Gaston and
Blackburn take the view that more careful and bal-
anced documentation of different patterns, including
negative findings, is needed. They list four approaches
to gain progress in macroecology: natural experiments,
microcosms, mathematical modeling and use of obser-
vational data.
Understandably, experimentation in the scale of in-
terest to macroecology is all but impossible and there-
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fore documentation of patterns before and after some
major changes in an ecosystem is a valuable source of
information. These changes can be caused by droughts
and other large-scale disturbances. Microcosms allow a
direct manipulation of a system under study, but the
choice of organisms is more limited. Modeling can help
in clarifying issues concerning the origins of different
patterns. However, this is a relatively undeveloped field
macroecology and thus provides much space for
innovation. A major problem with observational data is
formulating the null hypothesis, which is a clear indica-
tion of unripe understanding of the underlying pro-
cesses. Setting null hypotheses is important since they
help in separating trivial patterns from the ones imply-
ing a real biological process. Gaston and Blackburn
also observe one difficulty in differentiating amongst
competing hypotheses: some patterns may be genuinely
caused by multiple processes, perhaps operating at dif-
ferent scales.
Macroecology and scale
The scale problem is explicitly dealt with by Rosen-
zweig and Ziv (1999). What is the role of scale in
able predictions about local species diversity on the
basis of information on regional species diversity? One
way to pose the question is whether local assemblages
are random sub-assemblages of regionally available
species. Rosenzweig and Ziv argue that the theoretical
basis for this is to be found in species-area
relationships.
To answer the question what the scales in species-
area relationships have in common with the relation-
ship between regional and local diversity, we need first
to understand the mechanisms of species-area relation-
ships. To provide examples, extinction and speciation
are the mechanisms that govern diversity at the global
scale, and immigration and extinction determine the
species compositions on archipelago scales, or in meta-
populations. The local scales are determined by local
population dynamics. After understanding the mecha-
nisms of species-area relationships, we hopefully may
link these relationships to other ecological rules.
The question whether ecological rules (cf. Lawton's
conclude that the theoretical roots of the interpreta-
tions are not very substantial.
Huston (1999) addresses similar problems. He main-
tains that local and regional processes are intercon-
nected, so that local diversity can often be understood
only when knowing causes of regional diversity and
that regional diversity can only be understood via
processes acting on local diversity: "a biodiversity Yin
and Yang". The challenge for ecologists is to tell apart
87:3 (1999)
the relative importance of the two scales in affecting
each others' species richness.
Huston's argument is that local processes affecting
species richness, such as competition, can only be ex-
pected to show their effect under very strict conditions.
These "equilibrium" conditions are seldom met in na-
ture. Thus, Huston proposes that data on species num-
bers are not adequate for testing the relative
importance of local and regional effects. This is because
ecologists have rarely addressed the issue of species
richness on a proper local scale. For example, the data
taken from distribution range maps are very likely to
fail to fill strict criteria, because the spatial resolution is
too coarse and includes even at its finest scale a range
of different habitat types.
Despite the fact that climate affects large areas simi-
larly, Huston finds little evidence to attach ecological or
evolutionary processes uniquely regional in scale. In his
review, Huston is rather skeptical of the success of
contemporary ecologists in resolving the relative contri-
butions of local vs regional processes in providing an
understanding for global patterns of species diversity.
Scale and experimentation
The scale problem is also addressed by Petersen et al.
(1999). The past 50 years or so have witnessed how
experimentation has made ecology a more rigorous
science. The need for proper experimentation is
reflected by recent textbooks in experimental ecology
(e.g., Hairston 1989, Underwood 1997). It is challeng-
ing to find out which results from an experiment are
scale free and which are artifacts of the scale of the
experiment. Addressing questions of ecosystem func-
tioning with experiments would definitely call for differ-
of the largest unsolved issues of ecology. Against this
ent scale and temporal duration than unraveling
predator-prey interactions with bean weevils and their
parasitoids. The causes of the scale of enclosed aquatic
ecosystem experiments is the topic of the review by
Petersen et al. (1999).
Linking large-scale patterns to local processes
Stenseth (1999) advocates an optimistic stance. To-
gether, data and theory should be able to do the job of
disentangling general properties of ecological systems.
Few ecological features have captivated the mind of
ecologists more than the cyclic population fluctuations
of northern voles and lemmings (Elton 1924). Research
on vole and lemming ecology in Sweden, Norway,
Finland and Hokkaido has contributed much to the
contemporary understanding of patterns and processes
behind the cyclic dynamics of these herbivorous ro-
dents. Stenseth makes an effort to bring all the research
together. His review will certainly be the starting point
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for all who want to know where research on cyclic
rodent dynamics has led us during the 75 years since
Charles Elton brought the issue into daylight.
During the recent few years, Stenseth and his co-au-
thors have made a big effort to compile and analyze all
existing long-term time series on vole dynamics. The
analysis (using the most up-to-date statistical tools) of
periodically fluctuating small rodent population data
clearly suggests the presence of direct and delayed
annual density dependence. This implies that either a
predator-prey type of interaction or a plant-herbivore
type of interaction (but not both) may be the underly-
ing cause of the periodic multiannual population cycles.
The current consensus appears to be that specialist
predators are the key for understanding vole cycles.
However, clear-cut experimental evidence addressing
the validity of these propositions is still scanty (but see
Korpimaki and Norrdahl 1998).
In his review, Stenseth begins from the beginning
(Elton 1924) and goes through all major achievements
in the research to solve the puzzle of cyclic population
dynamics. One of the intellectual leaps forward is the
understanding that a given vole species may exhibit
different kinds of population dynamics in different
parts of its distribution range. This is clearly indicated
by the gradient of the multi-species vole cycle length
from southern Scandinavia (no cycle) to central (3
years) and the northern parts of this area (4-5-year
cycle). A similar gradient has been found in the gray-
sided vole data in Hokkaido. Together with the pattern
found earlier in Scandinavian vole data, this observa-
tion led to speculations of the significance of predators
behind the cyclic fluctuations. Nevertheless, the vole
and lemming cycle - despite the voluminous efforts of
trying to understand why the cycle occurs - is still one
background, it is worth mentioning that the enigmatic
vole cycle has started to disappear. In large areas of
Scandinavia the previously 3-5-year regular fluctua-
tions have leveled off (Henttonen et al. 1987, Hansson
1999).
Ecology in space
The ecology of spatially structured populations and
communities has a long tradition dating back at least to
Huffaker's (1958) classic experiments. As we have seen,
spatial scale plays a major role in macroecology. Land-
scape and metapopulation ecology are other ap-
proaches to the role of spatial structure. The only
anniversary review that explicitly deals with population
and community dynamics in space is Hanski's (1999)
metapopulation paper. Metapopulations are composed
of discrete habitat patches containing local populations
being woven together by migrating individuals. Local
extinctions and colonization of empty patches compose
(1999) 467
the dynamics of the metapopulation. Hanski derives
new measures to predict the dynamics of a focal species
using information about the structure of a fragmented
landscape. The aim is to develop a measure that could
be used to assess the persistence of a metapopulation in
a dynamically changing landscape. The novelty of this
idea is that the landscape where metapopulations are
living may often be in continuous change, new patches
being created and old ones permanently disappearing.
Thus, it may well be that part - if not all - population
turnover is due to turnover in the habitable patches
themselves. This habitat turnover may be natural, such
as successional stages replacing each other, or due to
anthropogenic causes. Hanski's new model (yes, in this
review there is genuinely original research) is a direct
continuation of his metapopulation work, and brings
the metapopulation concept from the classical Levins
(1969) metapopulation origins closer to contemporary
landscape ecology.
Evolution
Although Oikos is not specialized in evolutionary ecol-
ogy, it nevertheless reflects that strong back-bone of
modern ecology. Two of the anniversary reviews ad-
dress evolutionary problems.
Thompson's (1999) review deals with coevolution. As
so many other topics in evolution and ecology, the idea
now called coevolution was already presented by Dar-
win in The Origin of Species. For instance, in chapter 2,
he wrote: "...the structure of every organic being is
related, in the most essential yet often hidden manner,
to that of all the other organic beings...". Clearly,
Darwin saw the essence of coevolution in the interplay
between ecology and evolution. In his review, Thomp-
son focuses on mechanisms bringing about coevolution-
ary changes in nature and outlines the usage of
different kinds of data in coevolutionary studies. He
recognizes six main approaches in gaining understand-
ing of coevolution: studying (1) phylogenetic context,
(2) patterns of specialization, (3) genetic architecture of
coevolution, (4) distributed outcomes of interactions,
(5) rates and forms of selection, or (6) geographic
mosaic of interactions. These approaches enlighten dif-
ferent aspects of coevolution; some focus more on
constraints, others more on forms and possibilities of
coevolution. Thompson concludes that progress in this
field has suffered from lack of testable hypotheses,
especially due to the dominance of the paradigm of
diffuse coevolution. According to this idea, organisms
commonly interact with so many other species that
actual coevolution is rare or diffuse and therefore hard
to find. Intriguingly, this view can also be seen in the
Darwin quotation above when he wrote about the
hidden manners in the causation of structural related-
ness between organisms. Thompson argues powerfully
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against this logic. On the other hand, he warns about
making strong generalizations of the role of coevolu-
tion in community dynamics based on too narrow an
approach. He also claims that studies on coevolution-
ary processes are in a key position when evolutionary
studies are applied to community management. Possible
areas of applied research in coevolution include, for
instance, agriculture, conservation and epidemiology.
Every organism, in an evolutionary sense, represents
a life cycle. Therefore, when looking at a particular
trait and assessing its fitness value, one has to take into
account the whole life history of the organism. Due to
tradeoffs, a trait optimal at some life stage may be
suboptimal at another life stage. In his review,
Kozl'owski (1999) discusses the value of optimization
models in studies of adaptations. He emphasizes that
optimization models do not represent adaptationism;
they do not aim to prove that organisms are optimal,
but instead describe the optimization process of selec-
tion. Multiple reasons exist why the ultimate optimum
for a trait may never be reached. The difficulties in the
optimization approach include building a reasonable
model in the first place, transforming trait values into
fitness, the fact that model solution is not always an
ESS, and finally, it is not always clear how fitness
should be measured or defined.
The question about fitness is especially difficult to
solve. However, this is the issue that we are ultimately
interested in - and searching for an evolutionarily
optimal solution without a well-defined criterion is
doomed to fail. In addition, different fitness measures
often do actually affect the conclusions. Several alterna-
tives exist. It is known, for instance, that reproductive
value at birth is an appropriate fitness measure in
deterministic population dynamics. In real situations
this is a rather limiting assumption. Lifetime reproduc-
tive success (LRS) is a straightforward and often used
measure in empirical studies. However, there are
difficulties with LRS as well. Offspring may differ in
quality and under density dependence timing of repro-
duction matters to the spread of a genotype.
Mylius and Diekmann (1995) presented a solution to
density-dependent situations in Oikos. They asked, in
particular, which density-independent fitness measure is
maximized at an ESS. Alas, their solution depends on
the details of density dependence in the population
dynamics to such an extent that at the moment their
approach is not very practical with real data. Fortu-
nately, there are alternative ways to tackle fitness prob-
lems with optimization models as shown by Kozbowski.
None of these solutions is universal but will undoubt-
edly be useful in more specific situations. Kozlowski
also suggests ways to validate optimization models with
comparative methods. Further challenges for the future
include taking community structure and spatial dimen-
sion, e.g., metapopulation structure, into account in
optimization models.
87:3 (1999)
Genes in ecology
One of the scientific disciplines in natural sciences that
has experienced an exciting period of fast development
after the 1960s is molecular biology. Tunlid (1999)
reviews the recent development of molecular biology by
presenting it as a science linking microbial ecology,
general ecology and organismal biology. His main ar-
gument is that molecular techniques offer currently new
possibilities to identify species, population and commu-
nities of microorganisms in nature. Using the methods
provided by the recent development of molecular tech-
niques one can identify genotypes and species of mi-
croorganisms. In this way we can obtain information
on the structure of microbial communities, the evolu-
tionary relationship of the organisms, the spatial and
temporal dynamics of microbial populations, gene ex-
pressions and metabolic activities. In particular, it has
become possible to link microbial diversity to ecological
processes and develop the interaction between experi-
mental microbiology, microbial ecology, and general
ecology. Consequently, it should be able to narrow the
gap between ecology and other branches of biology.
Tunlid (1999) illustrates the great potential of molec-
ular biology in the tasks outlined above by presenting
examples of molecular methods that are used to reveal
the structures and functioning of microbial communi-
ties and populations in natural environments. Apart
from microbial communities, fungal DNA has been
sequenced from plant roots infected by pathogenic and
symbiotic (mycorrhizal) fungi. Molecular biology meth-
ods can also be used to analyze the spatial and tempo-
ral structures of microbial and other populations. For
example, bacteria in mixed species assemblages can
outgrow predation pressure, or in some other cases,
develop inedible, inactive morphological structures. The
recent development of genetic markers has made it
possible to test, for instance, the degrees of sexual and
asexual reproduction in hermaphrodites.
Ecology, environment and conservation biology
Conservation biology is a discipline taught at universi-
ties throughout the world. A huge number of people
work at institutes carrying the name conservation biol-
ogy in the academic world, in the private sector and in
public governmental positions. Conservation biology is
an important way for mankind to address the relation-
ship between economic, industrial and other activities
of the human society and the environment.
Despite the undeniably important role of conserva-
tion biology in all sectors of society, and despite the
fact that universities openly advertise conservation biol-
ogy as a new scientific discipline, the question of its
merit as a scientific discipline pops up regularly on the
pages of scientific journals. This is also the question
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that Noss (1999) addresses in his review: "Whether
conservation biology is a distinct discipline with its own
theories, methods, and applications - or simply an
amalgam of what scientists and practitioners in many
disciplines have been doing for many years - is a
troubling question to conservation biologists who yearn
for a disciplinary identity". He ends up evaluating this
problem by posing the question whether self-pro-
claimed conservation biologists are filling useful roles in
the scientific and conservation communities that have
not been filled by academic ecology and the resource
management disciplines. We may develop the theme by
asking whether applied ecology, applied genetics, or
resource management, or even environmental econom-
ics could do the same job. In practice, this issue has
developed as a professional rivalry between conserva-
tion biology and other disciplines such as wildlife biol-
ogy, applied ecology and environmental management.
Conservation biology appeared for the first time in
1937 on the pages of the Journal of Wildlife Manage-
ment (Errington and Hamerstrom 1937). At that time,
conservation was understood in North America as util-
itarian, as opposed to preservation, the latter referring
to protecting the wonders of Nature. Focus was on
game species of mammals and birds, as opposed to the
current approach of spreading out activities over all
taxa. The term "conservation biology" appeared regu-
larly in the late 1970s, with its content having changed
crucially. The beginning of conservation biology as a
unified discipline can be traced back to a symposium:
"Conservation biology: an evolutionary-ecological per-
spective" (Soul and Wilcox 1980). As is understand-
able, the idea must have been in the air for some time
then, as witnessed by many scattered journal articles
and several books published around the theme. The
term "biodiversity" developed as the guiding content of
conservation biology in the late 1980s.
In the 1990s, conservation biology has proceeded by
attempting to develop general principles for conserva-
tion and rules-of-thumb about population viability, re-
serve design, and similar issues. At the same time,
conservation biology has been closely tied to the tack-
ling of case studies. Consequently, it has been observed
that developing general laws in conservation biology is
jeopardized by the fact that all case studies seem to be
too specific.
As a set of selected examples of the principles devel-
oped in conservation biology we may consider the
following: 1) habitat in contiguous blocks is better than
fragmented habitat; 2) maintaining viable ecosystems is
usually more efficient, economical, and effective than a
species-by-species approach; 3) biodiversity is not dis-
tributed randomly or uniformly across landscapes; in
establishing protection priorities, consider hot spots.
The first principle states that habitat fragmentation is
usually deleterious for biodiversity. The second princi-
ple addresses the question of the object of conservation.
(1999) 469
Should we concentrate on protecting rare bird and
butterfly species, or should we take an alternative ap-
proach and protect functional entities in nature, such as
ecosystems? The third principle deals with the criteria
to characterize and compare different ecosystems or
landscapes. All these problems have high priorities in
conservation biology at the moment.
According to Noss, one of the key issues of contem-
porary conservation biology is whether its principles are
sufficiently general to guide conservation actions in
particular cases. Obviously, it is not a simple task to
manage natural populations or ecosystems. Fisheries
economics, for example, has been said to be a history of
mismanagement cases (Pitcher et al. 1998).
Another key issue raised by Noss for conservation
biology is the question of education and training. Most
of the conservation biologists have studied some tradi-
tional academic discipline such as biology or ecology,
and many of them have studied resource management,
forestry, or fisheries. However, it is not difficult nowa-
days to find university programs under the title of
conservation biology, alone or combined with wildlife,
fisheries, etc. We can distinguish at least two basic
problems in developing the education in conservation
biology.
The first deals with the multidisciplinary nature of
conservation biology. Biology and ecology are at the
heart of conservation biology training. Here we should
learn the basics of ecosystem functioning. The accom-
panying question is how far we can go with this under-
standing and how much we should be able to deal with
social sciences in order to successfully tackle conserva-
tion problems. Many scientists share, after all, the
opinion that conservation deals more with politics,
economy and other social issues than biology (see Haila
below).
The second problem deals with the relationship be-
tween empirical work and conceptual or theoretical
development. The rapid development of theory, both in
conservation biology and related sciences such as ecol-
ogy, population biology, and population genetics
makes it important to educate conservation biologists
to work in terms of general concepts and mathematical
models.
Habitat fragmentation and biodiversity
Harrison and Bruna (1999) focus their critical review
on habitat fragmentation and its consequences. They
correctly point out that habitat fragmentation in vari-
ous forms has been one of the occupations of ecologists
since the discipline of conservation biology began. Har-
rison and Bruna's argument is that the two extant
flagship theories, the theory of island biogeography
(MacArthur and Wilson 1967) and metapopulation the-
ory (Hanski and Gilpin 1997), do not, in fact, help us
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much to understand the ecological processes in frag-
mented landscapes. This is because the central element
in these theories is in dispersal linkage of mainland to
islands or population sub-units to each other. They
propose that these theories make too many simplifying
assumptions (no habitat quality differences among
patches, local population sizes reduced to presence/ab-
sence, focusing on non-interacting species, etc.) to cap-
ture the essence of fragmentation. They claim that "As
always, theory may either be an elegant simplification
that allows us to focus on the essential issues, or such
an oversimplification that its predictions are never met
in real systems because the factors it excludes are too
important".
Harrison and Bruna maintain that most of the extant
theory focuses on fragmentation as a spatial problem.
Thus, it is conjectured that dispersal of individuals in
the landscape has a central role in affecting the persis-
tence of local populations in a fragmented world. In
contrast with this view, Harrison and Bruna bring
evidence from empirical studies suggesting that it is the
quality of habitat that goes down with increasing frag-
mentation. This deterioration is due to the edge effect:
micro-climate changes, matrix species entering the
pristine habitat and edge-to-area ratio increases. Their
skeptical attitude is sobering reading and one cannot
but agree when they say that there can be few tasks
more important for ecologists than determining how
diversity can be maintained in the remnants of natural
habitat.
The unique versus the general is also dealt with by
Austin (1999). He is especially interested in the role of
vegetation ecology in biodiversity research. How should
we develop it? He defines plant community ecology as
the study of distribution and behaviour of multispecies
assemblages of plants in time and space. In comparison,
vegetation ecology applies to the branch of plant com-
munity ecology where observational analysis is the
dominant research method. In answering the
paramount question, Austin addresses the problems of
communication between paradigms used in species rich-
ness studies and asks whether vegetation ecology is able
to contribute to the study of biodiversity. In clarifying
the content of his review he raises three questions: 1)
The relevance of vegetation ecology's own paradigmatic
assumptions regarding theory, methodology, and exper-
imentation to current questions regarding biodiversity;
2) The need to simultaneously test hypotheses from
different paradigms in order to develop a new synthesis;
3) The use of vegetation survey and analysis techniques
in conservation biology.
Austin advocates the continuum as a pivotal but
neglected concept in vegetation ecology. Species com-
position and other collective properties such as species
richness vary as a continuum in a multi-dimensional
environmental space, and thus the concept or the pat-
tern of continuum should be included in the develop-
ment of the paradigms of biodiversity. He proceeds by
87:3 (1999)
posing the demand that any process-based ecological
theory should be able to take into account the contin-
uum in the sense that it should be able to predict the
ecological patterns, e.g. species richness, which occur
along environmental gradients. He ends up with em-
phasizing that both environmental space and geograph-
ical space need to be incorporated into the community
ecology paradigms.
Environmental gradients may or may not refer to
climatic ones. In addition, resource gradients can occur
and it is critical that these gradients are distinguished
when considering the continuum in space. Furthermore,
we must not forget the scale: depending on the geo-
graphical scale (global, regional or local) the patterns
may remain the same, indicating power law structures,
self-similarity, etc., or they may change, the patterns of
which remain largely unknown.
What are the reasons for the fact that the paradigms
used by vegetation ecologists are neglected by the ecol-
ogists in other branches? Austin refers to the general
lack of testable hypotheses as the major reason, and
lists a number of these. Obviously, a scientific discipline
which is not capable of generating ideas or theories that
could be verified by empirical tests is useless.
The second reason for the under-representation of
vegetation ecology in biodiversity research is, according
to Austin, methodology: the one-sided use of ordina-
tion methodology by many vegetation scientists. The
now-famous comment by May (1985) must have been
painful: "the wilderness of meticulous classification and
ordination of plant communities in which plant ecology bad and every human action that changes the environ-
has wandered so long, began in pursuit of answers to ment cannot by the statement above be evil and
questions but then became an activity simply of its own
sake".
Aliens
Invasions of alien species is a major conservation prob-
lem in many areas, especially in Australasia and the
USA. This is the topic of the review by Williamson
(1999). Questions for ecologists are along the lines of
"Why do some species become pests?" and "Can we
predict which areas are prone to invasion?". Under-
standing invasions is also extremely important for bio- Haila also argues that there is ultimately a "radical
control for two reasons. First, there is the pest
managers want to get rid off. Second, selecting a suit-
able control organism requires understanding the risk
of getting a new pest instead of successful biological
control. Apart from practical conservation and man-
agement, studying invasions sheds light on central eco-
logical issues. For instance, the question of the
relationship between ecosystem stability and species
richness has a long history. Invasion studies have
shown that in the tropics, invasion problems actually
tend to increase with biodiversity. However, very few
robust generalizations have been possible to make.
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High intrinsic rate of increase and absence of natural
enemies are typical to invading species but, for instance,
size, taxonomy and niche do not predict invasion prob-
ability successfully. Actually, the only consistent predic-
tor seems to be history: species having invaded
somewhere are likely to do it again.
One of the major problems with invasion studies is
the bias towards successful invasions. A way forward,
in Williamson's view, is to get more and better data.
There are not enough studies on natural population
dynamics of invasive species, let alone experimental
manipulations.
"Socioecologies"
The role of ecology in environmental issues and conser-
vation is not, however, unproblematic. "A basic fact of
human ecology is that human beings depend on natural
systems, but they cannot depend on such natural sys-
tems with which they have no interactions". This quo-
tation is from Haila's (1999) review of ecology as an
environmental science. Haila's point of departure is the
observation that environmentalists (including conserva-
tion biologists and environmentally concerned profes-
sional ecologists) tend to detach humanity from
"nature" and to regard the "environment" as a dualis-
tic contrast to human culture. This, Haila argues, is
fundamentally flawed. Human activities are not a priori
detrimental.
The modification of the environment in which hu-
mans live is a inevitable consequence of being a biolog-
ical organism and that is what humans are. A way out
of this predicament is, according to Haila, to embrace
this fact but also to be careful when defining humanity.
"Humanity" is not a well-defined and unequivocal en-
tity. "Humans" do not destroy rain forests or pollute
rivers or cause soil erosion, only certain segments of
humanity with a certain history and social and cultural
context do that, and it is different from place to place
and from time to time.
contextuality" in which we find ourselves. That is, our
(like any other organism's) existence depends on partic-
ular conditions here and now that have arisen due to a
chain of historical processes, some of them coinciden-
tal. This does not imply a laissez-faire mentality that
ignores responsibilities and respect, but instead an obli-
gation to appreciate the environment that actually
makes our existence possible. Hence Haila's title of the
review: "Socioecologies". Environmental problems are
real problems inasmuch as they threaten our existence
or well-being, but this threat can only be understood in
historical, social and cultural contexts.
(1999) 471
 Haila is also critical of the notion that the conflict by increased per capita survival during winter. This
between the productivity of human systems vs the argument has since long been used by hunters to ex-
productivity of ecological systems is a zero-sum game. plain why harvesting is not necessarily a bad thing from
First, this dualistic contrast is impossible in the first a population perspective. Standard models of popula-
place. Second, the so-called PAT rule cannot work. The tion and community dynamics tell us otherwise, but
PAT rule says that Impact = Population x Affluence x that is, Boyce et al. argue, because they do not account
Technology (Ehrlich and Ehrlich 1990). It cannot work for the seasonality. Should that be done, however,
because the right-hand terms are not independent, they models and a good body of data, mainly from birds
are not divisible to add up to given number, there are and mammals are often compatible. Boyce et al. show
no interaction terms, and they are not quantifiable in a that such models are relatively straightforward to for-
straightforward manner. It also omits the fact that the mulate and that both discrete time and continuous time
Earth is not a uniform whole and that humanity is not models lend themselves to do the job. A harvested
a unified entity. population characterized by sequential density depen-
Where does the science of ecology fit into all this? dences may actually have a higher equilibrium popula-
The annoying notion that real ecological principles and tion density than an unharvested one. Some of the
processes can only happen in pristine environments models that Boyce et al. review also make the predic-
should be abandoned. Ecology should then take on the tion that harvesting may affect the dynamics of the
role of characterizing the nature of Nature, human-infl-population. Low to moderate harvest rates seem to be
uenced and virgin habitats alike. The abandoning of the able to stabilize the dynamics whereas high harvest
balance of nature view is one such scientific break- rates do the reverse.
through according to Haila. Ecology should also These potential effects on the target population still
provide the raw material for action rather than just hold true if trophic-level interactions are included in the
reaction. In order to become operational, ecologists modeling. Although Boyce et al. only hint at the com-
must therefore learn how to make decisions (cf. Hilborn
plications (but also better understanding) that multi-
and Mangel 1997). Having given up the romantic view species models of harvested populations may invoke,
of Nature as an untouchable virgin and acknowledging they certainly have opened the road for ways of im-
the social, historical and cultural contexts of human proving population management by injecting harvesting
actions, as well as interactions with our environment, theory with some useful community ecology.
ecologists can start acting to make the Earth a better
place to be.

Whither ecology in the 2000s?

Population management
Conservation biology certainly also involves a great
deal of population management. Although relatively
little of that is expected to be found on the pages of
Oikos, but more on Ecography's, one of the anniver-
sary reviews deals with the topic.
Most ecosystems are characterized by seasonality. As
a result, reproduction often takes place during a limitedfuture, as seen by the authors of the anniversary re-
favorable time of the year. Other seasons are harsher
even for survival alone. Boyce et al. (1999) take this
fundamental observation seriously and argue that if
such sequences of events are associated with density-de-
pendent survival and/or reproduction, then it should
have notable consequences for population dynamics.
Such sequences of density-dependent processes should
also give room for compensation of predation or har-
vesting mortality.
Imagine, for example, that density-dependent repro-
duction happens in the spring. In the winter, there is
strong density-dependent mortality. Should this popu-
lation be harvested in the fall, before the yearly "bottle-
neck", then population density could be reduced to the
extent that per capita mortality rate during winter is
reduced. The loss due to harvesting is compensated for
Where shall ecology go, which are the future main
streets, where do we encounter new frontiers, and
which will be the battlefields of ecology? These are
intriguing questions and it takes a visionary to predict
what kind of ecology Oikos and her kind will promote
on their pages during the next few years to come, not to
mention perspectives as long as 50 years ahead. The
views, is listed in Table 1. Remember that the summary
should be read with caution since the authors of the
reviews were not asked to provide future perspectives.
Table 1 therefore represents our interpretation of what
we thought the authors felt would be important in the
future.
It should be obvious that a handful of reviews cannot
summarize the state of the art of ecology, which ever
since Huxley has developed from natural history to a
scientific discipline rich in subdivided fields. Many of
the topics that some ecologists would argue is the core
of ecology (e.g., plant-herbivore interactions, para-
sitoids and pathogens, ecosystems ecology, sexual selec-
tion, the list could be made infinitely long), even those
actively advanced by ecologists all over Fennoscandia,
were not touched upon in the anniversary reviews due

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Table 1. The "future of ecology" according to the authors of the Oikos-Ecography 50 years' reviews. This list is taken from the
corresponding reviews. Recall that in assembling this table we had to be terse due to space limitations. This list is put up simply
for giving insights into what the different authors felt the future should bring. Note that they were not advised to conclude their
essays with sweeping perspectives and conclusions. If the author(s) did not give indications of future the entry is left empty.

Austin
For Austin, continuum is a central concept to vegetation ecology. Consequently, he concludes that patterns, such as species
richness, dominance, standing biomass and assembly rules, should be detectable in relevant environmental spaces. He also
calls for the development of statistical modelling techniques for vegetation ecology.
Boyce et al.
Seasonality is a very important property of many, if not most, natural environments. However, this is seldom addressed in
population models. Boyce et al. show that seasonally structured models will be important for population management and
harvesting.

Brown
We need theory and models to understand emerging macroecological patterns and to formulate hypotheses causing these
patterns.

Gaston and Blackburn

Progress in macroecology will gain by more careful and balanced documentation of patterns and better understanding of
the mechanisms behind the patterns.

Haila
Haila warns against duality thinking: one could, and especially should, avoid culture-nature dualism in theoretical
developments as well as in practical aspects of conservation biology, wild life management, fisheries, etc. But most
importantly, even in desperate situations from the societal point of view, ecology might point out new possibilities to
overcome the problems which seem hopeless.

Hanski
It is important to notice that not only populations but also habitats can be dynamic. This is a way to greater unification
of landscape ecology and spatial population dynamics.

Harrison and Bruna

The most important task for ecologists is to determine how biodiversity can be preserved with ever-continuing
fragmentation of habitats. They call for empirically motivated development of theory that begins with real scenarios of
fragmentation, and has an explanation of the real observation as its goal.

Huston
Complains about the lack of data in testing ideas concerning species diversity. According to him, there are more ideas and
hypotheses around than data allow to test.

Kozlowski
Great progress in ecology has been because of an evolutionary approach. This direction will probably change. Evolutionary
studies will start gaining from acknowledging ecology in action. Populations do not live in density-independent
homogenous space. Acknowledging density dependence and spatial structure of populations is a great challenge to
evolutionary research for some time to come.

Lawton
Does not directly provide fuel for forthcoming studies, but rather just concludes that in ecology large enough scales
(macroecology) and small enough scales (individuals, populations) are quite well understood. "It is the middle ground
which is the mess".

Noss
A need for an integrative field of conservation science will remain for a long time to come. Conservation biology will
continue to make a positive contribution both in scientific and professional issues. He ends up with presenting a vision
where conservation biology will gradually become less distinct as its theories and techniques are incorporated into other
disciplines.

Persson
Disagrees with Polis. In his world, restricted subsamples ("community modules") should be studies as a caricature of real
ecosystems. He would like to see future ecosystem studies focusing on adaptive foraging, nutrient recycling, habitat
coupling and size-structured dynamics.
Petersen et al.
Stress the importance of scale in experiments. To be able to successfully extrapolate from lab to nature, physical and
temporal scales of the study have to match those of the organisms and systems studied.
Polis
The following questions need to be answered to understand ecosystem level distribution and dynamics of biomass: 1) How
does population dynamics affect ecosystem processes and vice versa? 2) How does spatial variability affect populations and
communities? 3) How does past productivity affect current interactions? 4) How do age- and stage-structured processes
affect food webs and communities? 5) What happens to the primary productivity not eaten by herbivores?
Rohde
Studying the deviations from the common pattern that species richness increases toward the equator should aid in
understanding latitudinal gradients in diversity. Also, appreciating that the patterns we see today often reflect
non-equilibrium situations will help our understanding.

OIKOS 87:3 (1999) 473

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Table 1. (Continued)

Rosenzweig and Ziv

Propose that community ecology will flourish from the realization that identifying the scale-dependent properties, in particu-
lar large scales, may help in revealing the determinants of communities. They also suggest that the study of assembly rules
has a promising future, and that it may be aided by expanding its analyses to include such topics as coevolution.

Stenseth
Takes, with wise caution, the effort to guide future research on cyclic population dynamics of voles and lemmings: 1)
Perform experimental manipulations for populations known to display multi-annual periodic cycles (trying to make them
only seasonal cycles) and for populations known to exhibit only seasonal density cycles (trying to make them multi-annual
cycles). Here lies the understanding of this enigmatic problem. 2) Integrate demographic analysis better into population
dynamic models.

Thompson
Studies on coevolutionary processes are in key position when evolutionary studies are applied to community management.
Possible areas of applied reseach in coevolution include, e.g., agriculture, conservation and epidemiology.

Tunlid
Asks what is the ecological significance of microbial diversity. He would also like to know what is the cause of microbial
diversity. Do nutrients in ecosystems cycle faster if microbial diversity is higher?

Williamson
Questions the predictability at invasions: it could be that invasions are just as unpredictable as earthquakes. Nevertheless,
despite this comparison, he maintains that better understanding of ecological invasions is possible, and such understanding
is badly needed in the context of preserving biodiversity.

to space limitations. Thus, the visions expressed in the
reviews, by necessity, remain limited in scope. However,
anyone who takes the time and effort to read all of
them will see that there still emerge topics in the
outlooks that many ecologists would agree are
important.
Common to almost all authors of the reviews is the
emphasis on the relevance of proper theory guiding the
empirical research. Thus, they echo the opinion held by
many others, most clearly put by Haila and Jarvinen
(1982): "Sound naturalism is to ecology what legs are
to a runner; but anti theoretical naturalists are, quite
naturally, like headless runners". Depending on the
author, and the broadness of issue dealt with in the
anniversary review, the theory called for is either a
sweeping one, or very detailed. We agree that at this
stage of the history of ecology as a scientific discipline
there still is a long way to the ecological Theory of
Everything. However, it is hard to believe that ecology
as a natural science will be a network of localized
theories rarely communicating across the (often arbi-
trary) boundaries of ecological sub-disciplines. Thus, in
this field there is plenty of free space for genuinely new
thinking, by ecologists as well as by philosophers.
Against this background, we suggest that the tour
through the 50th anniversary begins with Lawton's
is about time. During the past 75 years (since Elton
1924) ecologists have kept on debating with both theory
and data as arguments to give the answer to the causes
and effects of cyclic dynamics in voles, grouse, hare and
lynx.
To call for better data and better tools to deal with
the data does not come as a surprise. New methods
have always proved to provide more precise and de-
tailed answers that the old ones. One has to keep in
mind, however, that now and then ecologists also have
to rethink their questions. Ecology has seen several
"paradigm shifts" during the past 50 years, clearly
revealed when browsing through the pages of Oikos
from volume 1 to volume 87. One thing is certain in
ecology, that paradigms will continue to die and be
born and some will even become re-born. In this intel-
lectual stream the society will continue to challenge
ecologists to providing answers to various environmen-
tal issues be they due to habitat degradation, or loss of
biodiversity rising from the conflict between contrasting
needs of economy and concern of our well-being being
mediated by the ecological state of the Earth.
Mingled with the needs of the society, ecologists will
also make an effort to thrive as researchers advancing
their own scientific discipline. Here, certainly adhering

review. It bravely takes the intellectual effort to tie it all to principles of the theory of evolution by natural

together. There are others who like to guide ecologists
to the new millennium with critical but not acutely
skeptical advice (Persson 1999, Brown 1999). Stenseth
(1999), who took on the paramount work of summariz-
ing the results of the multi-annual cycles industry (espe-
cially Nordic voles and lemmings), is very precise in his
visions of the future of ecology - to understand cyclic
dynamics. If his outlooks are to come true, one of the
largest unsolved issues of ecology will become solved. It
selection they will continue to pave the road to success.
And although evolutionary ecology seems to be unbeat-
able, there is always "plus ultra" - more beyond!
Whether molecular genetics or ecosystems research will
be the main channel for ecology the coming 50 years
remains to be seen. We know one thing for sure,
though. Ecology is very rich in data and has a wealth in
theory and this makes it both useful, fun and intellectu-
ally challenging.

474 OIKOS 87:3 (1999)

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