2318 Population Dynamics | Metapopulation Models
Summary
Metacommunity models offer the most complete view of
the factors maintaining species diversity that has been
described to date. The metacommunity concept realisti-
cally describes how both local and regional forces can
contribute to species diversity, and how the structure of
local communities can be altered by immigration from
other communities within the region. To date, metacom-
munity models consist of four broad kinds: neutral, patch
dynamics, species sorting, and mass effects (or source–
sink). There is a variety of empirical evidence for species
sorting and mass effects models, whereas patch dynamics
appear less common, and there is no good example of a
system with neutral community dynamics. A metacom-
munity structure is likely to alter the composition of local
communities, food web structure, species abundances,
and the potential for evolution.
See also: Average Taxonomic Diversity and Distinctness;
Biodiversity; Community; Island Biogeography;
Metapopulation Models; Ordination.
Further Reading
Cadotte MW and Fukami T (2005) Dispersal, spatial scale, and species
diversity in a hierarchically structured experimental landscape.
Ecology Letters 8: 548–557.
Chase JM and Leibold MA (2003) Ecological Niches: Linking Classical and
Contemporary Approaches. Chicago: University of Chicago Press.
Chave J (2004) Neutral theory and community ecology. Ecology Letters
7: 241–253.
Metapopulation Models
I Hanski, University of Helsinki, Helsinki, Finland
ª 2008 Elsevier B.V. All rights reserved.
Introduction
Classification of Metapopulation Models
Four Kinds of Metapopulation Models
Models for Dynamic Landscapes
Metapopulation Models for Two or More Species
Introduction
Most landscapes are complex mosaics of many types of
habitat. From the viewpoint of a particular species living
in such a landscape, only some habitat types, called sui-
table habitat, provide the resources that are necessary for
population growth. The remaining landscape, often called
Chesson P (2000) Mechanisms of maintenance of species diversity.
Annual Review of Ecology and Systematics 31: 343–366.
Cottenie K (2005) Integrating environmental and spatial processes in
ecological communitydynamics. Ecology Letters 8: 1175–1182.
France KE and Duffy JE (2006) Diversity and dispersal interactively affect
predictability of ecosystem function. Nature 441: 1139–1143.
Fukami T (2005) Integrating internal and external dispersal in
metacommunity assembly: Preliminary theoretical analyses.
Ecological Research 20: 623–631.
Hairston NG, Ellner SP, Geber MA, Yoshida T, and Fox JA (2005) Rapid
evolution and the convergence of ecological and evolutionary time.
Ecology Letters 8: 1114–1127.
Holyoak M, Leibold MA, and Holt RD (eds.) (2005) Metacommunities:
Spatial Dynamics and Ecological Communities. Chicago, IL:
University of Chicago Press.
Hubbell SP (2001) The Unified Neutral Theory of Biodiversity and
Biogeography. Princeton, NJ: Princeton University Press.
Leibold MA, Holyoak M, Mouquet N, et al. (2004) The metacommunity
concept: A framework for multi-scale community ecology. Ecology
Letters 7: 601–613.
Loreau M, Mouquet N, and Gonzalez A (2003) Biodiversity as spatial
insurance in heterogeneous landscapes. Proceedings of the National
Academy of Sciences of the United States of America
100: 12765–12770.
Mathiessen B and Hillebrand H (2006) Dispersal frequency affects local
biomass production by controlling local diversity. Ecology Letters
9: 652–662.
McCann KS, Rasmussen JB, and Umbanhowar J (2005) The dynamics
of spatially coupled food webs. Ecology Letters 8: 513–523.
McGill BJ, Maurer BA, and Weiser MD (2006) Empirical evaluation of the
neutral theory. Ecology 87: 1411–1423.
Mouquet N and Loreau M (2002) Coexistence in metacommunities: The
regional similarity hypothesis. American Naturalist 159: 420–426.
Urban MC (2006) Maladaptation and mass effects in a metacommunity:
Consequences in species coexistence. American Naturalist
168: 28–40.
Urban MC and Skelly DK (2006) Evolving metacommunities: Toward an
evolutionary perspective on metacommunities. Ecology
87: 1616–1626.
Vellend M (2006) The consequences of genetic diversity in competitive
communities. Ecology 87: 304–311.
Application to Landscape Management and
Conservation
Summary
Further Reading
the (landscape) matrix, can only be traversed by migrating
individuals. Often the suitable habitat occurs in discrete
patches (also called habitat fragments). Individual habitat
patches may be occupied by a local population of the focal
species, but many patches are likely to be unoccupied at a
particular point in time, because a local population went
extinct in the past or the patch appeared in the landscape
 Population Dynamics
only recently due to, for example, successional changes.
The currently unoccupied patches may become colo-
nized in the future. The set of local populations
inhabiting the network of habitat patches is called a meta-
population. The local populations are typically connected
to other populations by some degree of migration, but
how strong this coupling is depends on the structure of
the landscape (how far apart the habitat patches are
located) as well as on the powers of migration of the
species.
Mathematical metapopulation models are used to
describe, analyze, and predict the dynamics of metapopu-
lations in fragmented landscapes. This article presents an
overview of different kinds of metapopulation models,
with an emphasis on spatially realistic models, which
can be applied to real metapopulations for the purposes
of research, management, and conservation.
Classification of Metapopulation Models
There is no sharp distinction between models for
single populations and metapopulations. Classic single-
population models assume that individuals interact
equally with all other individuals (panmictic population
structure). In the other extreme are classic metapopula-
tion models, such as the Levins model described in this
article, which assume a set of dynamically independent
local populations. But there are also intermediate mod-
els. A model involving the spatial coordinates of
individuals and limited spatial range of interactions
and movements can be viewed as a detailed population
model, or it can be considered as a metapopulation
model at the landscape level. Such individual-based
models for continuous space may exhibit spatial varia-
tion in habitat quality and thereby offer a very general
framework for population modeling, but their potential
to address relevant questions about metapopulation
ecology remains largely unstudied and they are not
covered in this article. Other models include a descrip-
tion of the genetic structure of the metapopulation, and
models may be used to analyze evolutionary processes
in metapopulations. These models too are beyond the
scope of this article, which is restricted to ecological
metapopulation models.
Table 1 Four types of stochasticity affecting metapopulation dynamics
Type of stochasticity Entity affected
Demographic Individuals in local populations
Environmental Individuals in local populations
Extinction–colonization Populations in metapopulations
Regional Populations in metapopulations
| Metapopulation Models 2319
Stochastic versus Deterministic Models
Metapopulation dynamics are influenced by four kinds of
stochasticity (Table 1): demographic and environmental
stochasticity affecting separately each local population,
and extinction–colonization and regional stochasticity
affecting the entire metapopulation. Notice that the latter
two forms of stochasticity are analogous to demographic
and environmental stochasticity. Metapopulation dynamics
are inherently stochastic, because population extinction
and colonization are stochastic events, and real metapopu-
lations are additionally affected by regional stochasticity.
Nonetheless, stochasticity is often ignored and instead a
deterministic model is constructed and analyzed. An impor-
tant difference between stochastic and deterministic models
is that the former predict a smaller or greater risk of
metapopulation extinction, whereas the latter predict that
for some parameter values the metapopulation has a posi-
tive equilibrium. However, for large metapopulations the
stochastic model settles for a long time to what appears a
stationary state, called a quasi-equilibrium. Thus for large
metapopulations the essential behavior of the metapopula-
tion is well captured by a deterministic model.
Discrete-Time versus Continuous-Time Models
Population models can be constructed by assuming dis-
continuous changes in population size at discrete time
intervals, for instance once a year, or continuous temporal
changes in population size. Though in reality the sizes of
all populations change continuously, the breeding seasons
may be so short that a discrete-time model is warranted,
particularly if population regulation influences only cer-
tain age classes. In discrete-time metapopulation models
with population turnover, extinctions and colonizations
occur at time intervals that agree with the life history of
the species, for example, colonizations follow the season
during which migration occurs.
Number of Populations and Description of
Landscape Structure
How many populations does the metapopulation consist
of? How is the landscape structure represented in the
model? Models may consider only two local populations
connected by migration, which represents a minimal
Correlation among entities
No
Yes
No
Yes
2320 Population Dynamics | Metapopulation Models
metapopulation but is nonetheless sufficient to address
some general questions about the role of migration in
local dynamics. In the other extreme, other models
assume an infinite number of habitat patches to simplify
model analysis; these models are typically focused on the
processes of population extinction and colonization. The
treatment of space can be implicit, explicit, or realistic. In
the first case, the model includes no information on the
spatial locations of habitat patches and local populations,
which implies that all local populations are equally
coupled to each other. The archetypal spatially explicit
model assumes a regular lattice, where lattice cells repre-
sent habitat patches. Finally, spatially realistic models
involve a description of habitat patches in terms of their
spatial coordinates, areas, qualities, and so forth, in a
manner that allows the application of the models to net-
works of real habitat patches.
Four Kinds of Metapopulation Models
Two-Population Models
The simplest metapopulation model extends models for
single populations to two populations coupled by migra-
tion. For instance, local dynamics may be modeled with
the familiar logistic model, and migration by assuming
that a fraction m of individuals migrates to the other
population. Migration may greatly influence the size of
the metapopulation, depending on the form of population
regulation, the difference in the carrying capacities in the
two patches, and mortality during migration. If local
dynamics are modeled with discrete-time models, which
are inherently less stable than continuous-time models,
more complex dynamics may emerge. Migration may
now stabilize local populations that exhibit complex
dynamics in the absence of migration, but migration
may also amplify population fluctuations, all depending
on the details of the model and the exact parameter
values. Even limited amount of migration may bring
population fluctuations into synchrony.
The two-population models may be extended to n
populations, but typically at the cost of the analysis
being restricted to simulations. n-population models
have been used to study metapopulation viability for
conservation. Such models are appealing to ecologists
and conservationists because of their apparent realism,
but because of the typically large number of untested
structural model assumptions and unmeasured parameter
values one cannot place much confidence on model
predictions.
The two-population and n-population models have
been used to study source-sink population dynamics.
Some interesting results include the possibility of sink
populations enhancing metapopulation stability when
source populations exhibit large fluctuations leading to
high risk of extinction. A metapopulation consisting of
independent sink populations with temporal variation in
growth rate may persist even if long-term growth rate is
negative in each population in the absence of migration.
Migration among such populations enhances metapopu-
lation growth rate by spreading the risk of locally bad
period among many independent populations.
Levins Model
The Levins model has a special significance for metapo-
pulation ecology, as it was with this model that Richard
Levins introduced the metapopulation concept in
two papers published in 1969 and 1970. The Levins
model represents a completely different approach to
metapopulation modeling in comparison with the two-
population model. The assumptions of the Levins model
are: (1) The suitable habitat occurs in infinitely many
patches that are equally large and of the same quality.
(2) The patches have only two possible states, occupied
versus empty, and hence the Levins model is an example
of patch occupancy metapopulations models, in which
local dynamics are ignored. (3) Local extinctions and
colonizations occur independently in different patches.
(4) All local populations are equally connected to other
populations and patches, which is another way of saying
that the model is spatially implicit.
With these assumptions, the size of the metapopula-
tion can be described by the fraction of the currently
occupied patches, denoted by p(t). Temporal changes in
the value of p(t) are given by
dpðt Þ=dt ¼ cpðt Þð1 – p½t Þ – epðt Þ ½1
where c and e are colonization and extinction rate
parameters. The first term describes the rate of coloniza-
tion of currently unoccupied patches (fraction 1p of
all patches), while the second term gives the rate of
extinction of currently occupied patches (fraction p).
Local extinctions may be due to demographic and envi-
ronmental stochasticity, but at the metapopulation level
local stochasticity translates to a constant extinction rate
parameter. Note that because the existing local popula-
tions are assumed to be identical, they all contribute
equally to the rate of colonization, and hence the coloni-
zation term is proportional to p as well as to 1p.
The equilibrium metapopulation size is given by
p ¼ 1 , where  ¼ e/c is called the extinction threshold.
In the Levins model a species that can invade a currently
unoccupied patch network has a positive equilibrium size
in the network, and vice versa (Figure 1a). This is not so
in more complex models that may have alternative stable
equilibria (see section titled ‘Population size-structured
models’). The basic reproductive number R0 in the Levins
model is given by 1/  , which has to exceed unity for the
species to be able to spread into an empty patch network.
 Population Dynamics | Metapopulation Models 2321

(a) (b)
1.0 1.0

0.9 0.9
Fraction of occupied patches, P

Fraction of occupied patches, P

0.8 0.8

0.7 0.7

0.6 0.6

0.5 0.5

0.4 0.4

0.3 0.3

0.2 0.2

0.1 0.1

0.0 0.0
–0.1 –0.1
0.0 0.1 0.2 0.3 0.4 0.0 0.1 0.2 0.3 0.4
Colonization rate Colonization rate

Figure 1 This figure shows the metapopulation equilibria in relation to increasing colonization rate. Locally stable equilibria are shown
by continuous line, unstable equilibria by broken line. (a) In the Levins model, the metapopulation may invade a patch network at the
extinction threshold, at which point metapopulation extinction (p ¼ 0) becomes an unstable equilibrium and only the positive
equilibrium is stable. (b) In population size-structured models with strong rescue effect, there are two alternative stable equilibria for
some parameter values, in which a metapopulation that is originally sufficiently large may persist in a network that it cannot invade from
small size (in the example colonization rate between 0.11 and 0.2). From Hanski I (1999) Metapopulation Ecology. Oxford: Oxford
University Press.

The Levins model is a deterministic approximation of 1000

a model known as the stochastic logistic model, which is
Number of patches n

an example of stochastic patch occupancy models. The

stochastic logistic model assumes a finite network of n 100
patches. If a patch is occupied, it is assumed to go extinct
with a fixed rate E ¼ e, while the colonization rate of an
empty patch is assumed to depend on the fraction of 10
occupied patches, C ¼ ck/n, where k is the number of
currently occupied patches. These assumptions define a
Markov process with metapopulation extinction as an 0.2 0.4 0.6 0.8 1
absorbing state, that is, sooner or later the entire metapo- Occupancy state p *
pulation will go extinct. A key prediction of the model is
Figure 2 The number of habitat patches needed to make the
the mean time to extinction, T. Using a diffusion approxi- mean time to metapopulation extinction at least 100 times longer
mation to analyze the model, which is well justified when than the mean time to local extinction. The dots show the exact
the number of patches is large, we obtain: result based on the stochastic logistic model, while the line is
rffiffiffiffiffi based on the approximation given by equation (2). From

2 e – ðn – 1Þp Ovaskainen O and Hanski I (2003) Extinction threshold in
T¼ ½2 metapopulation models. Annales Zoologici Fennici 40: 81–97.
n p2 ð1 – p Þn – 1

where p is the size of the metapopulation at quasi-equili-

brium. Figure 2 gives the number of patches n that the
network must have to make T at least 100 times as long as
the expected lifetime of a single local population. For
metapopulations with large p a modest network of
around ten patches is sufficient to allow long-term persis-
tence, but for rare species (say p < 0.2) a large network of
n > 100 is needed for long-term persistence.
The stochastic logistic model includes extinction–
colonization stochasticity but no regional stochasticity,
which leads to correlated extinctions and colonizations.
In the presence of regional stochasticity the mean time to
metapopulation extinction does not increase exponentially
with increasing n as predicted by eqn [2] but as a power
function of n, the power decreasing with increasing corre-
lation in extinction and colonization rates. This result is
analogous to the effects of demographic and environmental
stochasticities on the lifetime of single populations.
Population Size-Structured Models
The stochastic logistic model extends the Levins model
by incorporating extinction–colonization stochasticity in
2322 Population Dynamics | Metapopulation Models
the model. Population size-structured models represent
another type of extension. These models are deterministic
and retain the assumption of infinitely many identical and
equally connected patches, but the models include a
description of local dynamics and migration in the same
manner as the two-population models. The simplest
structured model divides existing local populations into
just two classes, small and large, which is sufficient to
model one important new process in comparison with
the Levins model: migration from existing large popula-
tions (which send out many migrants because they are
large) may increase the growth rate in small populations
and thereby rescue them from local extinction. At the
metapopulation level, the rescue effect leads to the
novel prediction of alternative stable equilibria, one of
which is metapopulation extinction, the other one a posi-
tive metapopulation size (Figure 1b). Therefore, a
metapopulation may persist in a network that cannot be
invaded from small metapopulation size.
Spatially Realistic Metapopulation Models
An important simplification of the models that have been
discussed so far is that they involve no description of the
landscape structure, hence it is not possible to apply the
models in a quantitative manner to real metapopulations.
Real habitat patch networks consist of a finite number of
patches that typically differ in their size and quality, and
in their spatial connectivities to existing populations,
which affect colonization rate.
To model metapopulations in such heterogeneous
patch networks, we extend the stochastic logistic model
to situations in which the extinction and colonization
rates (in continuous-time models) or probabilities (in
discrete-time models) are specific to particular habitat
patches. This leads to heterogeneous stochastic patch
occupancy models. Below, a deterministic approximation
of the full stochastic model is discussed, which leads to a
family of models with a clear relationship to the Levins
model and to models that are helpful for practical
applications.
In continuous time, the deterministic model is defined
by a set of n equations for a network of n patches,
dpi ðt Þ=dt ¼ Ci ð p½tÞð1 – pi ½t Þ – Ei ð p½tÞpi ðt Þ
where p is a vector of probabilities of the n patches being
occupied, pi, and Ci and Ei are patch-specific colonization
and extinction rates. To link colonization and extinction
rates to the structure of the fragmented landscape, we
make assumptions of how these rates depend on the
properties of the habitat patches, leading to spatially
realistic metapopulation theory.
Generally, the extinction risk decreases with increas-
ing patch area Ai, because large patches, when occupied,
tend to have large populations with a small risk of extinc-
tion. A reasonable parametric assumption is Ei ¼ e=Ai ex ,
where e and  ex are two parameters. The colonization rate
increases with connectivity to existing populations, with
connectivity defined as a measure of the expected rate of
migration from all possible source populations to the focal
habitat patch. A reasonable parametric assumption of
connectivity is
X  
Si ¼ A im
i exp – dij A em
j pj ½4
j 6¼1
Thus connectivity of patch i is obtained as a sum of
contributions from all other patches from which migrants
may arrive. The contribution of patch j increases with its
area Aj, scaled by power  em; with decreasing distance
between patches i and j, with 1/ giving the average
migration distance; and with the probability of occupancy
of patch j, as migrants can only arrive from occupied
patches. Immigration to patch i may depend on its own
area, scaled to power  im. The colonization rate is then
defined as Ci ¼ cSi.
The full dynamics of the model is described by the
system of n coupled equations [3], but its essential behav-
ior is well approximated by just one equation,
dp =dt ¼ c9p ð1 – p Þ – e9p ½5
Here metapopulation size is measured as the weighted
average P of the patch-specific occupancyPprobabilities
pi, p ¼ Vi pi, where the weights Vi ( Vi ¼ 1) are
derived from the relative contributions of individual
patches to the capacity of the landscape to support a
metapopulation. Large and well-connected patches have
larger values of Vi than small and poorly connected
patches, because large patches have populations with
small risk of extinction and send out many migrants to
colonize new patches, especially if they are located close
to the other patches. Note that this model is structurally
identical with the Levins model, though metapopulation
size is measured in a different manner and the coloniza-
tion and extinction parameters are defined as c9 ¼ cM/!
P
and e9 ¼ e/!, where ! ¼ Vi Ai. The new quantity here is
M, which is called the metapopulation capacity of the
fragmented landscape. Mathematically, M is the leading
eigenvalue of a ‘landscape’ matrix, which is constructed
½3 with assumptions about how habitat patch areas and con-
nectivities influence extinctions and colonizations. To
compute M for a particular landscape one needs to
know the scale of connectivity, set by parameter  in
eqn [4] for connectivity, the scaling of immigration, emi-
gration, and extinction rates by patch area ( im,  em, and
 ex), and the areas and spatial locations of the habitat
patches. The size of the metapopulation at equilibrium
is given by
p ¼ 1 – =M ½6
 Population Dynamics
An attractive feature of the spatially realistic models is
the possibility of estimating the values of model para-
meters with empirical data on the structure of the patch
network (patch areas, spatial coordinates, and possibly
other information) and spatiotemporal pattern of patch
occupancy. Data should preferably be available for sev-
eral years, including many extinction and colonization
events. Having estimated model parameters based on
data from a particular landscape, one may predict meta-
population dynamics and persistence in other landscapes,
for instance to guide management and conservation
actions. The models are most appropriate for highly frag-
mented landscapes, where all habitat patches are small or
relatively small and the respective populations have
hence a substantial risk of extinction.
Models for Dynamic Landscapes
The vast majority of metapopulation models assume that
the landscape structure remains unchanged, only the
population sizes and the spatial pattern of habitat occu-
pancy changes. This is a reasonable assumption for many
but not all species and landscapes. In particular, metapo-
pulation structures are common in many successional
habitats, in which changing habitat quality is an important
reason for local extinctions. Human-caused habitat loss
and fragmentation are often so fast that one cannot
assume metapopulations to occur at quasi-equilibrium
with respect to the current landscape structure, and we
may ask how long it takes for a metapopulation to reach
the new quasi-equilibrium (which may be metapopula-
tion extinction) following a change in landscape structure.
This latter question will be addressed in the section
‘‘Transient dynamics and extinction debt’’ below.
One may extend a stochastic patch occupancy model
to include turnover in habitat patches. A general formula-
tion for the stationary probability of occupancy of patch i
in a network of n patches is given by
Ci
Ji ¼ ½7
Ci þ Ei
Ji is often called the incidence of occupancy, and the
model an incidence function model. Assuming that exist-
ing patches disappear (which increases the extinction
probability of the respective local population) and new
patches appear in the landscape, the incidence function
may be written as
Ci – Ci ð1 – Ci – Ei Þage
Ji ¼ ½8
Ci þ Ei
where age is the age of the patch. Note that initially the
patch is always unoccupied (Ji ¼ 0 when age ¼ 0), while
the incidence in old patches (age large) approaches the
incidence given by eqn [7]. Given data on the ages of the
| Metapopulation Models 2323
patches in addition to data on patch areas and connectiv-
ities and the incidences of occupancy, this model may be
parametrized in the same manner as other stochastic
patch occupancy models. The size of the metapopulation
at quasi-equilibrium is given by the sum of the patch-
specific incidences. Given that Ei in eqn [8] is greater than
in eqn [7], it is clear that landscape dynamics reduce
metapopulation size.
Metapopulation Models for Two or More
Species
Species living in fragmented landscapes interact with other
species. Metapopulation models have been constructed for
competing species and for prey and predator inhabiting the
same patch network. This has led to several important
results. For instance, a species with a high colonization
rate may coexist at the landscape level with a species that
is a superior competitor locally, essentially because the
former species finds a temporary refuge in those patches
that have not yet been colonized by the superior compe-
titor. Similarly, a prey that is driven to extinction locally
may persist in a network of habitat patches if it has a
sufficiently high colonization rate. Spatially restricted
range of movements and ecological interactions may give
rise to strongly aggregated spatial distributions of interact-
ing species in the absence of any environmental
heterogeneity, which has been studied with continuous-
space and discrete-space (lattice) models. The latter are
conceptually related to metapopulation models, especially
if one assumes that lattice cells (representing habitat
patches) are occupied by local populations rather than by
single individuals. Spatial pattern formation due to ecolo-
gical interactions is less likely to be a dominant feature in
metapopulation models for fragmented landscapes, because
the fixed spatial variation in habitat quality strongly con-
strains (meta)population dynamics.
Application to Landscape Management
and Conservation
Habitat Loss and Extinction Threshold
Loss and fragmentation of natural habitats due to human
land use is the most important reason for the current
catastrophically high rate of loss of biodiversity on
Earth. Population viability depends, among other things,
on the environmental carrying capacity and hence on the
total amount of habitat available. Additionally, the spatial
configuration of habitat may influence viability, because
most species have limited migration ranges and hence not
all habitat in a highly fragmented landscape is readily
accessible. Metapopulation models have been used to
2324 Population Dynamics | Metapopulation Models
address the population dynamic consequences of habitat
loss and fragmentation.
In the Levins model, habitat loss has been modeled by
assuming that fraction 1  h of the habitat patches becomes
unsuitable for occupancy and reproduction, while fraction
h remains suitable. Habitat loss reduces the colonization
rate to cp(h  p), because the model assumes that fraction
1  h of the migrants land on unsuitable habitat and perish
(the model thus most literally applies to species with long-
range passive migration and no habitat selection). The
species persists in a patch network if h exceeds the thresh-
old value  set by the extinction-proneness and
colonization capacity of the species. An interesting impli-
cation of this result is that, at equilibrium, the fraction of
suitable but unoccupied patches (h  p) is constant and
equals the amount of habitat at the extinction threshold
(h ¼ ). This is a potentially helpful result, because it
suggests a way of measuring the value of the extinction
threshold given knowledge of h and p. The model is
however exceedingly simple and hence not really suitable
for quantitative predictions.
The spatially realistic metapopulation models com-
bine the metapopulation perspective of the Levins
model with a description of the spatial distribution of
habitat in a fragmented landscape and how the landscape
structure influences the extinction and colonization pro-
cesses. In the model described by eqn [5], the
metapopulation capacity M replaces the fraction of sui-
table patches h in the Levins model, and the threshold
condition for metapopulation persistence is accordingly
given by M > . The novelty here is that M takes into
account not only the amount of habitat in the landscape
but also how the remaining habitat is distributed among
the individual habitat patches and how the spatial config-
uration of habitat influences extinction and colonization
rates and hence metapopulation viability.
The metapopulation capacity can be computed for
multiple landscapes and their relative capacities to support
viable metapopulations can be compared even without
knowledge of the threshold value : the greater the value
of M, the better (Figure 3). One complication is regional
stochasticity, which can only be included in the calcula-
tions if there is empirical knowledge about the spatial scale
and the strength of stochasticity. Without regional stochas-
ticity, it always helps to have as short distances as possible
between habitat patches. With regional stochasticity, meta-
population viability is likely to be maximized by
intermediate distances between the habitat patches: long
enough to reduce the correlation in local dynamics, but not
too long to reduce migration too greatly.
Transient Dynamics and Extinction Debt
A change in the structure of a fragmented landscape, for
instance a reduction in the area of some habitat patches, will
1
0.8
0.6
p *λ
0.4
0.2
–3 –2.5 –2 –1.5 –1 –0.5 0
log10 Metapopulation capacity (λ M)
Figure 3 Metapopulation size of the Glanville fritillary butterfly
(Melitaea cinxia) as a function of the metapopulation capacity in
25 habitat patch networks. The vertical axis shows the size of the
butterfly metapopulation based on a survey of habitat patch
occupancy in 1 year. The empirical data have been fitted by a
spatically realistic model. The result provides a clear empirical
example of the extinction threshold. From Hanski I and
Ovaskainen O (2000) The metapopulation capacity of a
fragmented landscape. Nature 404: 755–758.
influence the patch-specific extinction and colonization
rates, but it takes time before the metapopulation has
reached a new quasi-equilibrium following environmental
change. The length of the transient period is longer when
the change in landscape structure is greater, when the rates
of extinction and colonization are lower, and when the new
quasi-equilibrium following environmental change is
located close to the extinction threshold. The latter result
has important implications for conservation. Species that
have become endangered due to recent changes in land-
scape structure are located, by definition, close to their
extinction threshold, and hence the length of the transient
period in their response to environmental change is pre-
dicted to be long. This means that we are likely to
underestimate the level of threat to endangered species,
because many of them do not occur at quasi-equilibrium
with respect to the present landscape structure but are only
slowly declining due to past habitat loss and fragmentation.
Considering a community of species, the term extinc-
tion debt refers to situations in which, following habitat
loss, the threshold condition is not met for some species,
but these species have not yet had time to go extinct.
More precisely, the extinction debt at a given point in
time is the number of extant species that are predicted to
go extinct in the future because the threshold condition is
not satisfied for them.
Reserve Selection
Setting aside a sufficient amount of habitat as reserves is
essential for conservation of biodiversity. Reserve selec-
tion should be made in such a manner that a given amount
of resources for conservation makes a maximal

contribution toward maintaining biodiversity. In the past,
making the optimal choice of reserves out of a larger
number of potential reserves was typically done by select-
ing reserves that together would include the largest
number of species, without any consideration for the
long-term viability of the species. More appropriately,
we should ask the question which choice of reserves
maintains the largest number of species to the future,
taking into account spatiotemporal dynamics of species
and possibly and preferably also predicted changes in
climate and land use. Metapopulation models can be
incorporated into analyses that aim at providing solutions
to such questions.
Summary
Metapopulations are assemblages of local populations
inhabiting networks of habitat patches in fragmented land-
scapes. The local populations are typically coupled by
more or less migration among the populations.
Metapopulation models are used to describe, analyze, and
predict the dynamics of metapopulations. Models have
been constructed to investigate the consequences of migra-
tion on the pattern and synchrony of local dynamics, the
processes that may lead to spatial pattern formation due to
ecological interactions, the persistence of metapopulations
in patch networks in a stochastic balance between local
extinctions and colonizations, and the response of metapo-
pulations to changing landscape structure. Metapopulation
models for highly fragmented landscapes predict an
Methane in the Atmosphere
S A Pegov, Russian Academy of Sciences, Moscow, Russia
ª 2008 Elsevier B.V. All rights reserved.
Increase in Methane Concentration
Methane Sources
Increase in Methane Concentration
Measured increases of methane concentration in the
Earth’s atmosphere began approximately 18 000 years
ago, and for the period of 10 000–12 000 years ago its
concentration rose from 350 up to 700 ppbv. Coming
nearer our time, 200–2000 years ago, methane in the
atmosphere changed from 700 to 800 ppbv. One hundred
years ago, methane concentration was already at
900 ppbv. During industrialization, methane emissions
Global Ecology | Methane in the Atmosphere 2325
extinction threshold, a critical amount and configuration
of habitat that is necessary for long-term metapopulation
persistence. The models thus predict that with increasing
habitat loss and fragmentation, species will go extinct
before all habitat has been lost. Spatially realistic metapo-
pulation models combine a description of landscape
structure with a model of the extinction and colonization
processes. These models can be parametrized with empiri-
cal data and applied to real metapopulations for the
purposes of research, management, and conservation.
See also: Colonization; Fitness Landscapes.
Further Reading
Hanski I (1999) Metapopulation Ecology. Oxford: Oxford University
Press.
Hanski I and Gaggiotti OE (eds.) (2004) Ecology, Genetics, and Evolution
of Metapopulations. Amsterdam: Elsevier.
Hanski I and Ovaskainen O (2000) The metapopulation capacity of a
fragmented landscape. Nature 404: 755–758.
Hanski I and Ovaskainen O (2002) Extinction debt at extinction
threshold. Conservation Biology 16: 666–673.
Levins R (1969) Some demographic and genetic consequences of
environmental heterogeneity for biological control. Bulleting of the
Entomological Society of America 15: 237–240.
Levins R (1970) Extinction. Lecture Notes in Mathematics 2: 75–107.
Ovaskainen O and Hanski I (2002) Transient dynamics in
metapopulation response to perturbation. Theoretical Population
Biology 61: 285–295.
Ovaskainen O and Hanski I (2003) Extinction threshold in
metapopulation models. Annales Zoologici Fennici 40: 81–97.
Verheyen K, Vellend M, Van Calster H, Peterken G, and Hermy M (2004)
Metapopulation dynamics in changing landscapes: A new spatially
realistic model for forest plants. Ecology 85: 3302–3312.
Conclusions
Further Reading
and atmospheric concentrations increased quickly up to
1500 ppbv. For example, measurements for 1978–88
showed that near-ground concentration increased from
1520 to 1690 ppbv. This growth rate slowed in the
late 1980s and practically stopped in 1991–92, believed
to be a result of the repair of Soviet gas pipelines.
Most recently, methane concentration is rising again.
Taking into account significant fluctuations of methane
concentration in the atmosphere measured in different
places of the globe and at various times, the average rate