10.

1 Islands and conservation

Our species has had a profound influence on the ecology and biodiversity of the planet, altering the
composition and functioning of ecosystems across the globe. Our actions in some respects represent
merely the latest driver of change, concentrated largely within one interglacial of a period, the
Quaternary (the past c.2 Ma), which has seen repeated dramatic fluctuations in climate. During this
time, individual species have experienced alternating episodes of expansion and of contraction and
fragmentation of ranges and have consequently been re-sorted into combinations that have often
differed radically from those found today (e.g. Bush 1994). There is nothing new about range
alterations.

Now, however, it is humans who are driving range shifts and extinctions. We have done this for a long
period of our history, and we are doing so in an accelerating fashion and on a global scale (Saunders
et al. 1991; Bush 1996). Whether we see particular changes in ecosystem properties as beneficial or
detrimental can depend as much on our cultural perspective as on the nature of the changes
themselves. However, there is no doubting the profound importance of the human agent in
biogeographical terms: not only in extinguishing species but, as significantly, by bombarding areas with
sets of species gathered up from far-distant regions of the globe, breaking bounds of isolation that
have lasted for many millions of years, and in the case of oceanic islands for their entire existence.

The accelerated extinction of species on a planetary scale is something that concerns all societies and
it should therefore be a high priority for natural scientists to study the processes involved, to document
the changes involved accurately, to develop predictive models, and to provide scientific guidance for
the conservation of biodiversity. This field of study is termed conservation biology, and it is an area of
research that has expanded greatly in the last 20 years: yet, there remain many uncertainties
concerning the big picture.

In illustration, we are currently unable to say to within an order of magnitude how many species we
share the planet with (Gaston 1991; Groombridge 1992). A second illustration is that although a large
proportion of land plants and animals are believed to be rainforest species, we cannot be sure how
large a proportion, nor how much tropical forest exists, nor with much precision, how fast these
ecosystems are being degraded of their biodiversity on a global scale (Whitmore and Sayer 1992;
Grainger 1993). We know there to be a crisis, but we cannot be sure of its precise magnitude.

Nature conservation as a social movement is not motivated solely by a concern to reduce global
species extinction but is also imbued with other values and goals. However, a full consideration of the
theoretical and operational principles of nature conservation and the long history of the conservation
movement are beyond our present remit. Our purpose in this final section of the book is to consider
two broad themes.

First, in the present chapter, we examine the application of ideas derived from island ecological
biogeography to conservation problems. This research programme is focused largely on the loss of
species resulting from the reduction in area and fragmentation of ‘natural’ habitats and is based on
the premise that island ecological rules can be applied to ‘islands’ of particular habitat types within
mainland landmasses. This work forms part of the emerging field of conservation biogeography
(Whittaker et al. 2005).
Second, we return in the final two chapters to a focus on real islands, examining the particular
conservation problems of oceanic islands and some of the management solutions that have been
developed. The rationale for focusing on such islands in an island biogeography book is obvious, but
this focus is also justified by the wider significance of islands as repositories of biodiversity and by the
extent to which island endemics feature in the lists of recently extinct and currently endangered
species.

10.2 Habitats as Islands

"... the abundance of publications on the theory of island biogeography and applications [of it] has
misled many scientists and conservationists into believing we now have a ready-to-wear guide to
natural area preservation . . ." (Shafer 1990, p. 102)

It is generally accepted amongst conservation biologists that the ongoing fragmentation and reduction
in area of natural habitats is a key driver of terrestrial species extinctions at local, regional, and global
levels (Whitmore and Sayer 1992). The remaining areas of more-or-less natural habitats are
increasingly becoming mere pockets within a sea of altered habitat; in short, they are habitat islands.
It was, therefore, to island biogeography that many scientists turned in the search for predictive
models of the implications of fragmentation and for guidance as to how best to safeguard diversity.

The implications of increasing insularity for conservation were recognized in the seminal works of
Preston (1962) and MacArthur and Wilson (1963, 1967), with Preston astutely observing that in the
long run species would be lost from wildlife preserves, for the reason that they constitute reduced
areas in isolation. MacArthur and Wilson’s dynamic (EMIB) model provided the theoretical basis to
develop this argument, allowing conservation scientists to explore the outcomes of differing
configurations of protected areas, assuming them to act as virtual islands in a ‘sea’ of radically altered
and thus essentially empty habitats.

One early manifestation of this research programme was the so-called SLOSS debate, which posed the
question ‘given the opportunity to put a fixed percentage of land into conservation use is it better to
opt for a Single Large Or Several Small reserves?’ Here the focus is on the entire regional assemblage
of species and how to maximize the carrying capacity of the protected area (i.e. habitat island) system.
We will come back to this debate a little later, as we have organized this discussion to begin with the
smallest units: populations of particular species.

The most basic question is: ‘how many individuals are enough to ensure the survival of a single isolated
(perhaps the last remaining) population? The resulting figure is termed the minimum viable
population. We will consider this first. Many species of conservation concern are not, however,
restricted to a single locality—or at least, not yet. Rather, their survival in a region might be dependent
on a network of habitat islands. Most of this book has been concerned with real islands in the sea.

Is it realistic to expect habitat islands within continents to behave according to the same principles as
real islands? Instead of the barrier of salt water, a forest habitat island might be separated from
another patch of forest by a landscape of meadows, hedgerows, and arable land. The implications for
species movements between patches might be radically different. As we have seen, however, this is
probably too simple a distinction and too simple a question. Scale of isolation is crucial, as well as the
nature of the intervening landscape filter. Mountain habitat islands within continents, but surrounded
by extensive arid areas, may be much more effectively isolated than real islands that happen to be
located within a few hundred metres of a mainland. Many isolates are actually sufficiently close to one
another that their populations are in effect linked: they form metapopulations, the second theoretical
element that we will consider.
When we have added these tools to our armory, we can then consider the whole-system implications
of fragmentation and how ‘island’ approaches have contributed to their understanding.

Over the past few decades there has been a shift in ecology from what Pickett et al. (1992) term the
balance of nature paradigm, which sees the world as a self-regulating, balanced system, towards the
flux of nature paradigm, which Pickett et al. (1992) believe provides a more realistic basis for
conservation planning and management. This framework does not deny the possibility of equilibrium,
but recognizes equilibrium as merely one special state. The flux of nature viewpoint stresses process
and context, the role of episodic events, and the openness of ecological systems.

Conventionally, the ‘island theory’ input to conservation debates has been based on the general
assumption that a ‘natural’ area is in a state of balance, or equilibrium. Humans then intervene to
remove a large proportion of the area from its natural state, leaving behind newly isolated fragments,
which are thus cast out of equilibrium. Subsequently, these fragments must shed species as the system
moves to a new equilibrium, determined by the area and isolation of each fragment. This way of
thinking is entirely consonant with the balance of nature paradigm.

By contrast, in the present volume, we have stressed the importance of considering physical
environmental factors in island biogeography, and have shown that ecological responses to
environmental forcing factors are often played out over too long a time frame for a tightly specified
dynamic equilibrium to be reached. Moreover, relatively few areas of the planet, if any, are pristine,
untouched by human hand, so most areas that we may be fragmenting already contain the imprint of
past land uses (e.g. cultivation and abandonment, hunting) (e.g. Willis et al. 2004). Hence, if the
starting assumptions concerning the prior equilibrial state of the system are flawed, some of the
theoretical island ecological effects may, in practice, be fairly weak. These points are addressed in the
present chapter in the applied, conservation setting. Hence we will consider the physical effects of
fragmentation as well as the biotic, and we will examine what sorts of factors in practice cause species
to be lost from fragments.

10.3 Minimum Viable Populations and Minimum Viable Areas

How many individuals are needed? What is the minimum viable population (MVP)? By this, we mean
the minimum size that will ensure the survival of that population unit, not just in the short term, but
in the long term. It is often defined more formally in terms such as the population size that provides a
95% probability of persistence for 100 or for 1000 years. Attempts to calculate the viability of single
populations, i.e. whether the population exceeds the minimum requirement, are termed population
viability analyses (PVA). Important elements that have to be considered in PVA include demographic
stochasticity, species traits, genetic erosion in small populations, and extreme events such as
hurricanes or fires.

Demographic stochasticity of initially small populations can lead to losses from a series of isolates
without a need to invoke any common factor such as predation or loss of fitness. However, where small
populations persist for a reasonable length of time, they may also lose genetic variability as they pass
through bottlenecks. They may then lack the genetic flexibility to cope with either the normal
fluctuations of the environment or an altered environment, and they may also accumulate deleterious
genes. In short, they lose fitness (Box 10.1).

It is generally held as axiomatic that an increase in inbreeding in small populations reduces fitness in
animals, although unambiguous demonstrations of the effect in natural populations are relatively
scarce (Madsen et al. 1996). The basic rule of conservation genetics is that the maximum tolerable rate
of inbreeding is 1% per generation. This translates into an effective population size of 50 to ensure
short-term fitness according to the calculations of I. A. Franklin in 1980 (reported in Shafer 1990).
However, there will still be a loss of genetic variation at such a population size, and Franklin
recommended 500 individuals to balance the loss of variation by gains through mutation. These
commonly cited values appear to derive respectively from data from animal breeders and bristle
number in Drosophila, and so should be regarded as merely first approximations (Fiedler and Jain
1992).

Lacy (1992) noted that crudely estimated MVPs for mammals based on data for inbreeding effects
from captive populations varied over several orders of magnitude, and that it would be premature to
generalize on MVPs in the wild on this basis.

Typically, only a proportion of the adult population participates in breeding, and it is these animals
that form the effective population size, which is often substantially smaller than the total population
size (Shafer 1990). A study of grizzly bears in the Yellowstone National Park showed that to prevent
inbreeding rates exceeding 1%, an overall population size of at least 220 rather than 50 animals was
required (Shafer 1990).

It is intuitively reasonable that the loss of a broad genetic base is likely to increase the probability of
extinction. However, we learned in an earlier chapter that putting a population through a bottleneck
of just a few individuals may not be as damaging to the genetic base as one might anticipate, providing
the population is allowed to expand again fairly quickly. In illustration, the great Indian rhino was
reduced within the Chitwan National Park in Nepal (one of its two main havens) to an effective
population of only 21–28 animals in the 1960s. In this case, the bottleneck was short. Rhinos have a
long generation time, and it has since recovered to about 400 individuals. Genetic variation within the
population is still remarkably high (Tudge 1991).

Nonetheless, the Drosophila studies reviewed in Chapter 7 showed that sustained or repeated
bottlenecks do lead to loss of heterozygosity. Hence, although you may be able to rescue some,
perhaps many, species from a bottleneck of short duration, you cannot assume that this will apply to
a sustained or repeated bottleneck. Just because a suite of threatened species is hanging on in their
reserves now, it doesn’t mean that the same level of biodiversity can be sustained for a long time in a
fragmented landscape.