Dynamics of Metapopulations: Habitat Destruction and Competitive Coexistence

Author(s): Sean Nee and Robert M. May

Source: Journal of Animal Ecology, Vol. 61, No. 1 (Feb., 1992), pp. 37-40
Published by: British Ecological Society
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Journal of Animal
Ecology 1992,
Dynamics of metapopulations: habitat destruction and
61, 37-40 competitive coexistence
SEAN NEE and ROBERT M. MAY
AFRC Unit of Ecology and Behaviour, Department of Zoology, South Parks Road, Oxford, OX1 3PS, UK

Summary
1. We take a previously studied model for two species - one of which is com-
petitively inferior - coexisting in a patchy environment, and examine the effects
of removing patches (that is, of decreasing the amount of available habitat).
2. Habitat destruction or patch removal reduces the number (and proportion) of
patches occupied by the superior competitor, but can result in an increase in the
total number of patches occupied by the inferior competitor (even though there
are fewer patches in total).
3. It has long been appreciated that disturbance and destruction of habitat can
create edge effects and other ecological changes favouring 'weedy' species. The
present study suggests that patch removal can by itself favour such species, even
in the absence of other, concomitant changes.
4. An implication relevant to conservation biology is that habitat loss can bring
about changes in community composition in remaining patches, even if such
patches themselves undergo no intrinsic changes whatsoever.
Key-words: competition, conservation biology, habitat loss, metapopulations.

Journal of Animal Ecology (1992) 61, 37-40

competitors. We discuss the implications of these

Introduction
Many species exist as metapopulations in a frag-
mented world, facing inevitable extinction in any
occupied patch, but persisting regionally by dis-
persal into unoccupied patches. It has long been
recognized that two species may coexist as meta-
populations even if one is competitively superior
within a patch, as long as the inferior competitor
either disperses more effectively or has a lower
patch extinction rate (Skellam 1951; Hutchinson
1951; Levins & Culver 1971; Horn & MacArthur
1972; Slatkin 1974; Hastings 1980; Hanski & Ranta
1983; Hanski 1983, 1987), or even in the absence of
these off-setting advantages if the superior competi-
tor's distribution is sufficiently clumped (Atkinson
& Shorrocks 1981, 1984; Shorrocks, 1990; Ives &
May 1985).
In this paper we analyse the effect of patch removal
on the regional abundances of two such species. We
find that patch thinning decreases both the number
of patches occupied by the superior competitor and
the number of empty patches, i.e. patches occupied
by neither species. At the same time, the number
(and, of course, the proportion) of patches occu-
pied by the inferior competitor initially increases.
Thus, patch removal or habitat destruction can
actually increase the regional abundance of inferior
37
results for understanding biodiversity changes in a
changing world.
Model
For simplicity, we make the extreme assumption
that the inferior competitor, B, is unable to invade
a patch occupied by the superior competitor, A.
Furthermore, if A invades a patch occupied by B
then B is immediately extinguished. The analysis
can clearly be extended to less extreme cases where
the difference in competitive ability between the two
species is at some intermediate level (with results
that are qualitatively similar); we choose not to
pursue such elaborations. A and B have coloniz-
ation rates CA and CB respectively, and extinction
rates eA and eB. We will consider an environment
consisting of a large number of patches, of which
a fraction h (for 'habitable') is suitable for occupancy
by the species, although some may actually be empty
at any particular time. If we let x, y and z denote the
proportion of empty patches, A-only patches and
B-only patches respectively in the environment,
the following equations correspond to the kinetic
diagram in Fig. 1.
dx
- = -cAxy + eAy - CBXZ+ eBZ, eqn la
dt
38 dy_ 3. lowers the number of empty (but not destroyed)
Habitat dt -
Axy
-
eAy
+
CAZY, eqn lb patches.
destruction If the inferior competitor is a superior colonizer,
dz
then the overall number of occupied patches falls.
dt CBZX
- eBz -
CAZY. eqn lc
If it is an inferior colonizer, then the overall num-
This is actually a two-dimensional system because ber of occupied patches rises. When h falls below
x +y + z = h. a critical value of its extinction/colonization ratio
Subject to the feasibility constraint that x, y and z (h < eAlcA), the superior competitor can no longer
are all greater than zero, these equations have the persist. Beyond this point, only the inferior com-
equilibrium solution petitor is found and the number of patches it occu-
pies declines as h decreases further. Finally, as patch
1 removal or destruction continues (h decreases still
x* = (hcA-eA + eB) eqn 2a
CB further), the inferior competitor also disappears
(for h < eBIcB). Fig. 2 illustrates these numbers (or
eA
y*=
h- eAeqn 2b frequencies, expressed in relation to the pristine
CA number of patches) as functions of h, for the special
eA(CA + CB) eB hCA
case in which both species have the same extinction
z* = - - .eqn 2c rate, but the inferior competitor has a higher colon-
CACB CB CB
ization rate. Fig. 3 explains the result.
In this system, feasible equilibria are locally stable
and species with feasible equilibria can increase
when rare.

CAYX
Empty CA/ * ? Superior
patches(x) 0.. 4 competitor
eAy patches (y)

eBz -------- --~~.

CBZ CAZy

C eeI
CB CA
Inferior
competitor
patches (z) Fig. 2. Illustrating the relative proportions of empty(....),
species A-only ( ) and species B-only (- - -)
Fig. 1. The structureof the model analysedin the text. patches (x*, y*, z*, respectively) as functions of the frac-
See text for an explanationof the symbols. tion of all patches (h) that are habitable or have not been
destroyed. The figure shows the stable equilibrium sol-
utions of equations 1 in the simplified case when extinc-
A necessary condition for the inferior competitor tion rates are equal (eA = eB 3 e). For 1> h > e/cA, both
species coexist in the proportions given by equation 2; for
to persist in this system is e/cA> h > eIB, only the more vagile but competitively
inferior species B can persist (z* = h - eICB, x* = eICB);
CB CA eqn 3 and for h < eIcB all patchesare empty.
eB eA

This may be satisfied if the inferior competitor has

a fugitive life style characterized by a high coloniz-
ation rate: our typical notion of a 'weed'. But the Superior
inferior competitor may also persist if it has a lower competitor

colonization rate, provided it has an even lower

extinction rate as well. This case corresponds to,
among other things, a species which uses its local
resources in a frugal fashion and may fall victim to a
profligate glutton of depletable resources.
We can immediately see from the equilibrium sol-
utions that patch removal, i.e. lowering h, actually:
1. increases the total number of patches occupied
by the inferior competitor (i.e. increases the pro-
portion of all patches, destroyed plus still existing,
occupied by this inferior competitor);
2. lowers the number of patches occupied by the
superior competitor; and
Patch
removal
o Inferior
competitor
Fig. 3. The direct effect of patch removal on both species
is negative, lowering the number of new patches they
colonize per unit time. The direct effect of the superior
competitor on the inferior is negative in two ways: first,
it raises its overall extinction rate and, secondly, lowers
the number of patches available for colonization by the
inferior competitor. So patch removal indirectly benefits
the inferior competitor by lowering the frequency of
patches occupied by the superior competitor.
39
Discussion
S. Nee &
R.M. May As Lande (1987) has emphasized in a single species
context, we do not have to destroy all patches to
extinguish a metapopulation that persists by virtue
of a balance between local extinctions and recoloniz-
ations in a mosaic environment. Vaccination pro-
grammes that eradicate infection without having
to reach every individual host (or 'patch') provide
concrete illustrations of this principle (Anderson
& May 1985). The present note goes beyond earlier
work to emphasize the counter-intuitive effects that
patch removal can have on populations of compet-
ing species. Inferior competitors persist by virtue
of colonizing empty patches, so it is surprising that
removing patches increases their regional abundance.
It is often observed that habitat fragmentation or
patch destruction can favour 'weedy' species over
those that are more commonly of concern to conser-
vationists, and this phenomenon is usually attributed
to disturbance and/or edge effects favouring such
weedy species. While these conventional expla-
nations undoubtedly explain much of what is ob-
served, the present work emphasizes that patch
removal per se can favour weedy species and ex-
tinguish other species even in the total absence of
edge-effects or other forms of habitat disturbance
within the remaining habitats.
Weedy species are not the only kind to be favoured
by patch removal. Inferior competitors may be or-
ganisms with lower colonization rates but which
consume local resources in a slow, more sustainable
fashion and can be starved by overconsuming weedy
species. Focusing only on competitive relationships
of this sort, our results show that patch removal can
actually increase the biodiversity in the remaining
patches, as the number of patches that are occupied
by neither species declines. Obviously, this result
should not induce complacency about the conse-
quences of the habitat removal that is occurring
around the world. Rather, it serves to highlight the
fact that the consequences of patch removal for
biodiversity may be non-obvious. Understanding
these consequences is a challenging goal for pure, as
well as applied, research in ecology.
The notion that environmental degradation may
have counter-intuitive consequences receives fur-
ther support from a theoretical analysis of Hastings
(1980). His study explores the effects of changes in
patch extinction rates (e, in Model 1) on the number
of co-existing competitors. In a model in which all
the species in a competitive hierarchy have the same
patch extinction rate, e, Hastings finds that as e in-
creases, the number of species that coexist does not
necessarily change in any simple fashion, but can
rise and fall several times.
One study that is explicitly relevant to our work is
of the patterns of occurence of Daphnia species in
rock pools on islands with different numbers of
pools (Hanski & Ranta 1983; for a re-analysis, see
Bengtsson 1991). They showed that coexistence
between competing species which differ in competi-
tive and dispersive ability is facilitated by a larger
number of patches. This foreshadows our work,
the purpose of which is to provide simple, quali-
tative insights into the effects of patch removal on
equilibrium species abundances. Such questions are
enjoying a surge of interest in today's green intel-
lectual climate and are readily addressed with this
sort of phenomenological modelling (MacArthur
& Wilson 1967; Levins 1969). We will explore the
more complex consequences of patch removal on
predator-prey relationships elsewhere.
The model can be looked at from viewpoints other
than those adopted here. The consequences of patch
addition may be of more relevance to European
ecology, if the farmland that is being removed from
agriculture over the next decades is not simply
paved over.
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Received 25 February 1991; revision received 13 May 1991