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doi:10.1006.bulm.2000.0208
Bulletin of Mathematical Biology (2002) 64, 209–212
Book Review
Metapopulation Ecology, by Ilkka Hanski, 1999. Oxford Series in Ecology and
Evolution, Oxford University Press, Oxford, U.K. $45 (paperback), 313 pp.
ISBN: 0-19-854066-3.
Early experiments with herbivorous and predatory mites with a patchy distribution
of oranges on a tray provided the first ingredients to the today’s well-developed
areas of spatial and metapopulation ecology. Those experiments by Huffaker were
done in the early 1950s and revealed a number of fundamental features related
with the stability and patchiness of field populations (Huffaker, 1958). Using a
relatively simple experimental setup, Huffaker was able to show that the wild time
fluctuations of preys and predators were sustained under appropriate manipulations
of the available feeding area. This was done by covering some oranges with paper
and paraffin or by intermingling rubber balls of similar size. Barriers were used to
restrict migration of predators thus introducing spatial heterogeneity.
Under spatial patchiness with non-equivalent patches (oranges) the population
oscillations where shown to be stabilized instead of displaying the rapid self-anihi-
lation of prey and predators observed in the homogeneous system after only one
cycle.
These experiments in fact provided the adequate experimental test for early the-
oretical studies of Nicholson who already asked the question of how population
dynamics would operate in a non-homogeneous, instead of well-mixed environ-
ment (Nicholson, 1933). In this sense, Nicholson’s prediction of global persistence
of locally weakly dependent populations received a first confirmation. These ques-
tions were later developed by Levins using simple models of metapopulations (i.e.,
a ‘population of populations’ in Levins’s words) where a group of several local
populations are linked by immigration and emigration (Levins, 1970).
Levins’s contribution had a very important impact in further developments of
ecological theory and defined the basis for the current theory of metapopulation
dynamics.
Today a huge number of theoretical and field studies have shown the importance
and implications of the metapopulation concept and its relevance for management
of endangered species. Not surprisingly, this theoretical approximation is espe-
cially important for the rapid process of habitat destruction and fragmentation dis-
played by natural ecosystems. In this sense, Ilkka Hanski’s book Metapopulation
Ecology is a timely, broad and extremely useful contribution to the literature on
metapopulations, well suited for theoreticians and empiricists alike.
Hanski’s book involves a clear presentation of metapopulation models and avail-
able field data organized as follows: (a) mathematical and simulation models (Chap-
ters 2–7), (b) field studies (Chapters 8–10) and (c) a case study of the metapopula-
0092-8240/02/010209 + 04 $35.00/0 c
2002 Society for Mathematical Biology
210 Book Review
tion ecology of a single species of butterfly (the Glanville fritillary, Melitaea cinxia)
which is in fact a long-term study conducted by Hanski and other researchers in
Finland. Through all the chapters, that particular example is repeatedly used as an
illustration of a real example from field data. Unfortunately, the apparent lack of
other works compiling a good theoretical model, sampling design and statistical
analysis in a long-term research project makes this system the only fully complete
study in metapopulation research.
The prologue already provides an excellent overview of the area and how the
different approaches (one, two-population models, lattice models, etc.) are linked.
Theory starts with a neat presentation of classic, fundamental population biology,
always presented together with real examples as a reference. The two-population
metapopulation problem (the simplest in this context) gives special emphasis on
some key issues as the presence of complex dynamical patterns (such as spa-
tial chaos) and the source–sink population structure. An introduction to classi-
cal metapopulation research introduces many non-trivial aspects of metapopulation
patterns and their consequences (such as the rescue effect). An especially interest-
ing analysis is the median time to metapopulation extinction provided by stochastic
dynamics and its dependence on the number of habitat patches.
Spatially explicit approaches [which have been extensively explored over the last
decade, see Bascompte and Sol´
e (1995) and references cited] are dealt with by dif-
ferent types of models, based on cellular automata approximations, state transition
models and incidence functions, presented together with a section on parameter
estimation. Metapopulation genetics and interacting metapopulations are hot areas
of research, the first being largely introductory but certainly providing the most
interesting aspects of the problem and a good list of references for further reading.
The second includes some of the most interesting and counterintuitive results of
the recent literature on the effects of habitat destruction in multispecies communi-
ties, such as the so-called extinction debt (Tilman et al., 1994; Stone, 1995) and
the different scales affecting metacommunity dynamics.
Field studies are described in three parts dealing with spatial structure of popu-
lations, the prediction of species occurrence/absence of species in particular sites
and the implications of metapopulation theory for conservation issues.
The list of topics explored in these three chapters is impressive and it goes from
general problems to specific ones. An example of the first is the discussion of to
what extent is habitat loss the relevant factor instead of habitat fragmentation.
An example of the second would be the well-known study of the northern spotted
owl Strix occidentalis caurina) as a particularly interesting case study which played
a particularly relevant role in drawing the attention of conservation policies towards
spatial ecology.
Finally, several applications are discussed on the best known work of Hanski’s
group on incidence functions and includes the beautiful predictions of alternative
equilibria in the Glanville fritillary and the simple, but insightful example of the
American pika (Ochotona princeps).
Book Review 211
As Hanski points out, its Incidence Function Model (IFM) is applicable mostly
in highly fragmented populations, and much work has still to be done if some
prediction is to be achieved for metacommunities. However, the implications for
conservation policies and experimental and field designs are straightforward. The
most serious drawback, that of the assumption of quasi-stationarity for parameteri-
zation of the IFM, claims for more research on regional non-equilibrium situations.
In this sense, other purposes resting on the interplay between patch models and per-
colation theory (Keitt et al., 1997) might be an alternative basis of future progress.
In summary, the long-lasting need for a synthesis in ecology, makes an important
step forward with Hanski’s contribution. Applicability without lack of generality.
One’s impression: you can breath reality from theory.
REFERENCES
Bascompte, J. and R. V. Sol´
e (1995). Rethinking complexity: modelling spatiotemporal
dynamics in ecology. Trends Ecol. Evol. 10, 361–366.
Huffaker, C. B. (1958). Experimental studies on predation: dispersion factors and
predator–prey oscillations. Hilgardia 27, 343–383.
Keitt, T. H., D. L. Urban and B. T. Milne (1997). Detecting critical scales on fragmented
landscapes. Conserv. Ecol. (online) 1(1), 4. Available on the internet.
URL:http://www.consecol.org/vol1/is1/art4
Levins, R. (1970). Extinction. Lecture Notes in Mathematics 2, 75–107.
Nicholson, A. J. (1933). The balance of animal populations. J. Anim. Ecol. 2, 132–178.
Stone, L. (1995). Biodiversity and habitat destruction: a comparative study of model forest
and coral reef ecosystems. Proc. R. Soc. London B 261, 381–388.
Tilman, D., R. M. May, C. L. Lehman and M. Nowak (1994). Habitat destruction and the
extinction debt. Nature 371, 65–66.
RICARD V. SOLE,
Complex Systems Research Group,
Department of Physics FEN,
Universitat Polit`
ecnica de Catalunya,
Campus Nord, M`
odul B4-B5,
08034 Barcelona,
Spain
and
Santa Fe Institute,
Hyde Park Road 1399,
Santa Fe, NM 85701,
U.S.A.
212 Book Review
JAVIER G. P. GAMARRA,
Complex Systems Research Group,
Department of Physics FEN,
Universitat Polit`
ecnica de Catalunya,
Campus Nord, M`
odul B4-B5,
08034 Barcelona,
Spain
