EQUILIBLIUM OF ISLAND

The Theory of Island Biogeography

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The Theory of Island Biogeography

Cover of the first edition

Robert MacArthur
Authors
Edward O. Wilson
Illustrator John Kyrk
Country United States
Language English
Subject Insular biogeography
Publisher Princeton University Press
Publication date 1967
Media type Print (Hardcover and Paperback)
Pages 203
ISBN 0691088365

The Theory of Island Biogeography is a 1967 book by Robert MacArthur and Edward O.
Wilson.[1] It is widely regarded as a seminal piece in island biogeography and ecology. The
Princeton University Press reprinted the book in 2001 as a part of the "Princeton Landmarks in
Biology" series.[1] The book popularized the theory that insular biota maintain a dynamic
equilibrium between immigration and extinction rates. The book also popularized the concepts
and terminology of r/K selection theory.[2]

Contents
 1 Background
 2 Summary
o 2.1 Introduction
o 2.2 Area-diversity pattern
o 2.3 Modeling colonization and dispersion
 3 Field testing
 4 Impact
o 4.1 Applications
o 4.2 Criticism
 o 4.3 Legacy
 5 See also
 6 References
 7 External links

Background
The Theory of Island Biogeography has its roots in Wilson's work on the ants of Melanesia.
MacArthur synthesized Wilson's ideas about competition, colonization and equilibrium into a
simple graphical representation of immigration and extinction curves, from which one can
determine the equilibrial species number on an island.[3] MacArthur and Wilson's ideas were first
presented in a paper published in 1963,[4] and were further developed into a book.[3]

Summary
Introduction

In the introductory chapter, MacArthur and Wilson highlight the significance of studying island
biogeography. Since islands are less complex and more numerous than larger ecosystems, islands
provide better opportunities to develop insights and perform replicable field research. Given that
insular microcosms are common to all ecosystems, principles from island biogeography can be
applied generally.

This graph portrays the immigration rate of new species and the extinction rate of resident
species versus the number of species present on an island. The intersection point gives the
equilibrial species number.

Area-diversity pattern

In Chapters 2 and 3, MacArthur and Wilson postulate that insular species richness depends on
island size and isolation from source regions. The authors present an equilibrium model that is
based on the following concept: when there is an addition of the number of species on an island,
the island's immigration rate of new species will decrease while the extinction rate of resident
species will increase. MacArthur and Wilson thus assume that there will be an equilibrial point
where the immigration rate equals the extinction rate. They further hypothesize that an increase
in island size will lower extinction curves while a decrease in distance between the island and the
source region will raise immigration curves. Since the intersection of immigration and extinction
rate curves determines the species number, the authors predict that larger islands will have more
species than smaller islands (assuming these islands are comparably isolated) and isolated islands
will have fewer species than islands more proximal to source regions (assuming these islands are
equally large). There is additional discourse on how insular clusters and stepping stones affect
this model.

Modeling colonization and dispersion

Chapter 4 discusses survivorship theory. The authors describe a model which states that the
probability for successful colonization is dependent on birth rate, death rate, and carrying
capacity of the environment. From this model, conclusions are made on the average survival time
for a propagule's offspring, the average survival time of a saturated population, and
characteristics of successful propagules.

In Chapter 5, MacArthur and Wilson examine why species can be excluded from insular
environments and how the niche of a species changes after introduction. The authors surmise that
pioneering species can be excluded for the following reasons: the island has saturated levels of
pre-existing competition, the pioneering species cannot maintain a population large enough to
avoid extinction, and the island hosts too many or too few natural predators. When a species
colonizes a new area, the authors state that the species will either shift, expand or contract its
realized niche.

Chapter 6 is a theoretical exploration of dispersal models. The authors consider how insular
stepping stones affect the dispersion of species—particularly, the effects that size and isolation of
stepping stones have on dispersion. Further consideration is given to how dispersal curves and
average distance travelled by pioneers impacts this study.

In Chapter 7, the authors state that there are generally 3 consecutive phases to the evolution of
populations after colonization. Initially, there is a trend for colonizers to evolve from r-strategists
into K-strategists. The founder effect may also influence colonizing populations during this first
phase. The second phase is marked by long term adaptations to the local environment. In this
period, abilities for dispersal are commonly reduced, and colonizers will either differentiate or
assimilate with competing species. In the third phase, the evolution of colonizing populations
may result in speciation and/or adaptive radiation.

Field testing
The insular biota equilibrium theory was experimentally tested by E. O. Wilson and his then-
graduate student Daniel Simberloff in six small mangrove islands in the Florida Keys. The
islands were fumigated to clear the arthropod populations. Immigration of species onto the island
was then observed in a first and second year census. Wilson and Simberloff confirmed that there
was an inverse relationship between the number of species on an island and the distance to the
source region as predicted in The Theory of Island Biogeography.[5]

Impact
Applications

MacArthur and Wilson's theory of island biogeography has been widely applied outside of island
ecosystems. For microbiota, the theory has been applied to the distribution of ectomycorrihzal
fungi on trees,[6] the distribution of bacteria in water-filled treeholes,[7] and the distribution of
fungi among shrubs.[8] While for flora and fauna, the theory's predictions have been realized with
the species richness of plants on mountains[9] and with the species richness of aquatic snails in
bodies of water.[10] Novel applications looked at plants as islands for insect species[11] and the
dependence of the species richness of mites on the areas of the host ranges of rodent species.[12]
MacArthur and Wilson's work has been used as a basis in other ecological theories, notably the
unified neutral theory of biodiversity,[13] and has been foundational for the fields of landscape
ecology, invasion biology, and conservation biology.[3][1]

Criticism

Several studies have disputed the underlying assumptions in MacArthur and Wilson's theory of
island biogeography: specifically, the interchangeability of species and islands, the independence
between immigration and extinction, and the insignificance of non-equilibrial processes.[14] In the
2001 preface, Wilson stated that "the flaws of the book lie in its oversimplification and
incompleteness".[1]

Legacy

In 2007, a symposium was held at Harvard University honoring the fortieth anniversary of The
Theory of Island Biogeography.[3] Following this conference, a collection of papers was
published in the book The Theory of Island Biogeography Revisited.[3]

See also
 Landscape ecology
 R/K selection theory
 Conservation biology
 Unified neutral theory of biodiversity

References
1.

 MacArthur RH, Wilson EO (2001). The theory of island biogeography. Princeton, N.J:
Princeton University Press. ISBN 978-0-691-08836-5. OCLC  45202069.
  Pianka ER (November 1970). "On r- and K-Selection". The American Naturalist. 104
(940): 592–597. doi:10.1086/282697.
  Wilson EO (2010). "Island Biogeography in the 1960s". In Losos JB, Ricklefs RE. The
Theory of Island Biogeography Revisited. Princeton University Press. pp. 1–12.
ISBN 9780691136530. JSTOR  j.ctt7s5m6.6.
  MacArthur RH, Wilson EO (1963-12-01). "An Equilibrium Theory of Insular
Zoogeography". Evolution. 17 (4): 373–387. doi:10.1111/j.1558-5646.1963.tb03295.x.
  Simberloff DS, Wilson EO (1970). "Experimental zoogeography of islands. A two‐year
record of colonization". Ecology. 51 (5): 934–937.
  Glassman SI, Lubetkin KC, Chung JA, Bruns TD (February 2017). "The theory of island
biogeography applies to ectomycorrhizal fungi in subalpine tree "islands" at a fine scale".
Ecosphere. 8 (2): e01677. doi:10.1002/ecs2.1677.
  Bell T, Ager D, Song JI, Newman JA, Thompson IP, Lilley AK, van der Gast CJ (June
2005). "Larger islands house more bacterial taxa". Science. 308 (5730): 1884.
doi:10.1126/science.1111318. PMID  15976296.
  Belisle M, Peay KG, Fukami T (May 2012). "Flowers as islands: spatial distribution of
nectar-inhabiting microfungi among plants of Mimulus aurantiacus, a hummingbird-pollinated
shrub". Microbial Ecology. 63 (4): 711–8. doi:10.1007/s00248-011-9975-8. PMC 4108428.
PMID  22080257.
  Riebesell JF (May 1982). "Arctic-Alpine Plants on Mountaintops: Agreement with Island
Biogeography Theory". The American Naturalist. 119 (5): 657–674. doi:10.1086/283941.
  Lassen HH (March 1975). "The diversity of freshwater snails in view of the equilibrium
theory of island biogeography". Oecologia. 19 (1): 1–8. doi:10.1007/bf00377585.
PMID  28308826.
  Opler PA (1974). Oaks as evolutionary islands for leaf-mining insects. American
Scientist. 62: 67-73.
  Dritschilo W, Cornell H, Nafus D, O'Connor B (1975). Insular biogeography: of mice
and mites. Science. 190 (4213):467-9.
  Hubbell SP (2001). The unified neutral theory of biodiversity and biogeography.
Princeton: Princeton University Press. ISBN 9781400837526. OCLC 733057107.

14.  Brown JH, Lomolino MV (January 2000). "Concluding remarks: historical

perspective and the future of island biogeography theory". Global Ecology and
Biogeography. 9 (1): 87–92. doi:10.1046/j.1365-2699.2000.00186.x. ISSN 1466-822X.

Island biogeography has been a subject of considerable interest to biologists and

geographers since the time of Darwin, Wallace, and the less well-known Hooker.
Hooker explored islands in the South Atlantic and South Pacific. Darwin and Wallace
are more important in our current thinking, since these two were pioneers in the
development of the theory of evolution. However, much of the theory of island
biogeography was built on data which came from their studies of the Galapagos and
East Indies respectively. Islands have been studied as natural experiments ever since,
with varying levels of intensity. Oceanic islands are isolated and small enough to reveal
processes and results too complex to interpret in mainland areas.

Islands are unique. Since they are isolated, evolutionary processes work at
different rates - there is little or no gene flow to dilute the effects of selection and
mutation. Endemism is rampant. However, both in theory and practice, that same
isolation makes islands more vulnerable to habitat change and extinction. Introduction
of a single predator or herbivore can have dramatic impact on the local community, as
we have just seen with biota of the Hawaiian islands.

Islands, as natural experiments, have not been protected from damage and
extinction through human activities. Since islands are isolated, and in many cases the
species found on them are endemic, extinction has been particularly common on
islands. 93% of the bird species whose extinction has been recorded since 1600 have
been island species. Historical records suggest a mean extinction rate through the
Pleistocene of approximately 1 species in 83.3 years. In 1980, that rate was one
species every 3.6 years. Extrapolating the curve, an extinction rate of one island
species per year can be predicted for the immediate future.

One of the reasons islands are important in the more general structure of ecology,
biogeography, and conservation biology is that islands, as at least relatively isolated
areas, are excellent natural laboratories to study the relationship between area and
species diversity. When we fully understand the relationship, it will be applicable to
fragments of habitat that human activities protect. We all know those sanctuaries are
important, but we need to know what and how much we can protect in them.

Species-Area Relationships

One of the immediately obvious characteristics of islands is the number of species

resident there, a number much lower and therefore more countable than the diversity on
a continental mainland. Such counts can reveal interesting relationships. For example,
Great Britain has 44 species of mammals, yet Ireland, only approximately 20 miles
further removed from mainland Europe into the Atlantic, has only 22 species. Is 20 miles
a sufficient distance to increase isolation and decrease mammalian immigration by half?
If so, then flying mammals (bats) should show similar numbers of species on both
islands, since immigration and isolation would be significantly less severe to a bat
species. The number of bats is not similar. Only 7 of the 13 bat species resident in
Great Britain breed in Ireland. What factor accounts for the difference? The single factor
which provides the best explanation is island area (though it is not the only contributing
factor). There is a classic curve originally drawn by Darlington, and reprinted in every
biogeography chapter since then, which depicts the relationship between area and the
number of reptilian species in the West Indies.

Figure - A curve relating island area to reptilian diversity in the West Indies.

As a log-log plot it is not a 'curve', but a straight line. From it a 'rule-of-thumb' can be
formulated which says that increasing the area of an island by a factor of 10 (or, more
correctly, counting the number of species on a nearby island 10 times larger) would
approximately double the number of reptile species present. That relationship is called a
power function. For islands it is written as:

S = C Az

where:

S is the number of species present,

C is a constant which varies with the taxonomic group under study (taxa which consist
of good dispersers (these species also typically have rapid population growth) will
logically accumulate more species on an isolated island, all else being equal),

A is the area of the island, and the exponent z has been shown to be fairly constant for
most island situations.

Geographic variation in C has been observed and 'loosely' reflects the isolation of island
groups typically studied due, for example, to the presence or absence of major air or
water circulation pathways nearby (islands located along the Gulf Stream would be
more likely to accumulate species than those located in the doldrums of the Sargasso
Sea) and also to effects of gross climatic difference (i.e. C is higher in the tropics than
for islands at high arctic latitudes). C can also be regarded as a scaling factor. In terms
of the graph of a species-area relationship, z determines the 'shape' of the curve, which
is raised or lowered as a whole by the value of C. The meaning of z, in an all out
treatment, is related to the distribution of abundances of species, i.e. to the number of
species expected if the total number of individuals increases (as it would on a larger
island) and those species follow a Preston log-normal distribution of abundance (see
May 1975 for a full treatment). However, there is a simpler level of meaning for the z,
revealed by manipulating

the equation. The graph of the basic equation would show an exponential rise in the
number of species as area increased. If we take the logarithms of both sides of the
equation, we get something that should look easier to work with:

log S = log C + zlogA

 

If we conveniently forget that the equation has logs as its terms, then you can see that
this is the equation for a line, with the basic formula:

Y = b + mx

So that if we plot the log of the number of species against the log of island area,
we get a straight line, and the slope of the line is the coefficient z. The z, establishes the
characteristic relationship between the number of species and area for all taxa. It is a
general characteristic, and fittingly, it is related to the most basic hypotheses about the
way species are put together in communities. There is variation in z, but it reflects, in
theory at least, general patterns of isolation involved in the studies. Study of taxa in
mainland 'islands' less isolated from their surroundings will give a lower z value (though
much higher total species numbers) than would parallel studies of the same taxa on
oceanic islands. It is argued this results from species that could not live indefinitely
within the 'island' being observed as transients in mainland habitat 'islands', though they
would never be observed on oceanic islands. Is z really as independent of autecological
characteristics of species and effects of real world disturbance as this view might lead
you to think? The answer, at least from modeling studies to be considered below, is a
clear no. The value of z seems to vary with the growth characteristics of species and
with the frequency and intensity of disturbance (Villa et al. 1992).

Species-area curves have been generated for a large variety of places and taxa,
and the range of z values is remarkably small. The data in the table below fit fairly
closely with what would be predicted based upon the theory of species abundance
distributions. May (1975) gave an extended treatment of the mathematical relationships
among observed and theoretical abundance and diversity patterns. Here the treatment
will be limited to a more compact form (derived from a chapter on patterns in multi-
species communities in May (1976)). The most commonly observed distribution of the
relative abundance of species is a log normal. This means that a plot of the logs of
species abundance against the number of species (i.e. a histogram) follows a bell
shape. Sampled distributions usually have what is called a 'veil line'.

Figure - Lognormal abundance distribution for moth samples.

 
 Figure - The theoretical Preston log-normal species abundance distribution

The curves indicate the presence of a few common species (the right hand end of the
curve) and a larger number of species of intermediate abundance. We don't usually see
the left hand end of the curve (the very rare species) because we rarely sample enough
individuals to capture even one. A larger total sample moves the veil line to the left,
taking in more of the total curve. Preston used other data, a log-normal species
abundance distribution for birds in Maryland, to develop the log normal.

Theoretically, the Preston log-normal the z value should be 0.263. May says that the
problem is that z doesn't really tell us much, since within the constraints as S varies
from 20 to 10,000 species, the z value changes only from 0.29 to 0.13. It really turns out
that z is largely a mathematical property of the log-normal, and doesn't tell us much
about communities. If they are log-normal, then z will be at least close to .263.
Nevertheless, there is still some interest in patterns in z. To indicate that briefly, here's a
short table of some species-area relationships:

Group Location value

_____________________________________________________________

beetles West Indies 0.34

reptiles and West Indies 0.30

amphibians

birds North America-Great 0.17

Basin

Birds West Indies 0.24

land vertebrates Lake Michigan 0.24

Islands
 ants Melanesia 0.30

birds East Indies 0.28

land plants Galapagos 0.33

dipterans Cincinnati parks 0.24

mammals North America 0.1-0.2

The values of z for mainland areas are clearly lower, e.g. both diptera in parklands
and mammals over North America generally, than the values for isolated islands. As
mentioned above, the explanation for these lower values is the inclusion of transients in
species counts from small 'islands'. Consider a badger, with a home range of 200
square km, or a wolf with a home range of 400 square km, or even larger areas for
seasonal migrants like caribou or large predatory birds. These animals could easily be
included in counts from 'island' areas much smaller than their home ranges, i.e. they
could be present in a 'patch' of mainland area considered as an 'island' transients. They
could not survive in such small areas if isolated by significant barriers, but sampling
larger areas more comparable to their home ranges would then not add them to species
lists as if new; they are already on the species list from being present as transients in
the smaller areas. Oceanic islands do not contain such transients. Thus we get
measurements of species numbers which are 'too high', include transients, in small
mainland sample areas, and too few additions as larger areas are sampled. The slope
of the graph relating the number of species to island area, i.e. the z, is lowered. The
theory underlying the log normal and island biogeography implicitly assumes that all
species counted could be resident permanently within the 'island' area, this may not be
the case with some mainland species. Compare the slopes for bird species numbers on
islands in the East and West Indies, both >0.2, with that for boreal forest birds in the
Great Basin of western North America (0.16). All three values are for habitat islands, but
on mainland the slope is lower, and the likely reason is the presence of some 'transient'
birds in species counts within small habitat islands in the Great Basin. That may also
explain the low value for mammals, but what about the low value for Cincinnati flies?
Are they likely to be transients? The explanation here more likely lies in the assumption
that islands are truly isolated; immigration onto an island requires a jump dispersal, i.e.
the intervening habitat is assumed to be inhospitable. The same sampling artifacts that
led to low values for mammals do also apply here. Models assume island populations
are numerically self-sustaining, and have increasing probabilities of extinction on the
island as they become rarer. Other green areas, garbage, etc. may sustain fly
populations (and even permit reproduction!) in habitats between park 'islands' in the
urban area, permitting high rates of re-immigration, so that populations which might not
otherwise be sustained in a park remain. That 'rescue' is much less likely for oceanic
island populations.
 

Figure - Species-area curves for ants on New Guinea (relatively, a mainland) and the
isolated islands nearby. The island curve is steeper (a higher z) than the New Guinea
curve, as explained above.

Is the relationship between species and area linear?

There are theoretical reasons to expect a z of 0.263. However, in doing so we are

accepting that species-area curves are linear. There are some points to consider:

 When small areas are sampled, species packing limits observed richness. There
is only so much space, so much variation in resources
 When large areas are sampled, samples may incorporate different habitats,
communities and species.

Given apparent differences in what you might expect, is the linear equation predicting
z logical? There is at least one study that questions the assumption of linearity (Crawley
and Harral, 2001). They sampled Berkshire and the East Berks in England in nested,
contiguous quadrats, at 100 and 25km2, also 456 km2 contiguous quadrats and
replicates (not contiguous) from 0.01 m2 to 110 ha in Silwood Park. Initial results,
averaging the values obtained, don't differ significantly from the theory. They found
z=0.302 for samples from the Silwood research estate, and z= 0.267 for samples over
the larger area of Berkshire and all of Great Britain.

An important point about effects of area on species diversity may have slid by here.
The biological question is why does area affect species numbers? There are two
schools of thought:

1) The Preston canonical log normal distribution can be used to suggest that area
determines the total population size of the collection of species living there. Area
is the direct determinant of diversity, since the multiplicity of factors which
determine relative abundance and species diversity are prescribed, and
independent of the specific island of area being studied.

 
 2) The alternative school suggests that area is of only indirect importance. Area fits
because area is a good indicator of the amount of habitat diversity present on an
island. It is really the 'number of niches' that determines the number of species,
but there is no established method for counting, or even estimating the number
of niches in an environment. Instead, physical variables are usually measured.
The assumption is made that the two measures should be highly correlated.

Let’s look at some of the basic data used to verify the basic relationship between
species numbers and island areas. One of the frequent approaches is the use of
multiple regression models to determine what factors account for differences in species
diversity on islands. Jared Diamond (1973; Diamond and Mayr 1976) used this method
to study the distribution of bird species on New Guinea and its satellite islands. The data
were first fit to the basic equation; for these bird data the equation then reads:

S = 15.1 A22

for the satellite islands near New Guinea, and this relationship accounted for 81% of
the variation in the number of bird species among islands studied. When Diamond
instead measured the numbers of species in montane habitat islands on New Guinea
proper, only the constant changed, to 12.3. Figure 4 shows the numbers of species and
the locations of montane habitat islands. The central 'line' of New Guinea is a mountain
backbone, and forms the largest single island. Diamond discerned that a part of the
effect of increasing area was due to an increase in the maximum elevation observed on
larger (even habitat) islands, and the concommitant differences in the number of
habitats resulting from a greater range of elevations, which cause differences in climate.

Figure - A map of the montane areas of New Guinea and the diversity of montane birds
in those areas. Variation in species numbers is correlated with variation in altitude.

In addition to the number of species accounted for by area alone, each 1000m of
elevation 'caused' an increase of 2.7% in bird species diversity on New Guinea satellite
islands (and 8.9% in New Guinea habitat islands) on average. After the entry of this
term, the regression equation reads:

 
 S = 15.1 (1 + .027L/1000) A22

Diamond realized that the number of species on islands also reflected the
degree to which the islands were isolated by distance from source populations. The
more distant from the New Guinea source of species, the smaller the number of species
when islands similar in size and elevation were compared. The decrease is
approximately exponential with distance, and the number decreases by half for each
2600 Km on average. Thus, Pitcairn Island (the place where the Bounty's mutineers
ended up) is about 8000 Km from New Guinea, and has about 1/8 or 12.5% of the bird
species numbers on a similar area of New Guinea. When this factor in incorporated into
the multiple regression equation, the result is:

S = 15.1 (1 + .027L/1000) (e-D/3750) A22

The following figures show you how some of the data look, and why the distance
relationship is exponential. Figure 5 shows the basic species-area relationship for the
satellite islands around New Guinea.

Figure - Species-area relationship for birds of oceanic islands around New Guinea.
Differences in isolation are evident both in absolute numbers of species and in different
slopes for islands in different distance ranges.

Figure compares some of the data for satellite islands to the curve for areas of the New
Guinea mainland (called a saturation curve). The slope is clearly steeper for the islands
than for mainland areas of similar habitats. The open circles are points for islands
relatively near New Guinea, the filled circles for more distant islands. More distant
islands have fewer species for the same area.

Figure looks at that relationship. Each island diversity is assessed as the fraction of the
number of species expected in a mainland habitat of the same area. This % saturation
is plotted against distance from New Guinea.

 
 Other data sets have been analyzed using the same basic approach, but one can
significantly add to our understanding of the biology underlying island diversity patterns.
In a study of bird species diversity on the Channel Islands off Santa Barbara, California,
Power (1972) worked backwards from the physical variables to the biological variables
which correlated most closely with bird species diversity, using a method called path
coefficient analysis. The method begins with a multiple regression analysis paralleling
the study of New Guinea satellite islands. Power first found the factor which explained
the largest portion of the total variation in bird species diversity, entered it into the model
equation, then determined the factor which explained the largest portion of the variation
which remained after applying the model equation. That factor enters second, and forms
part of a new model equation. Subsequent factors are entered in order of importance,
using the variation about the model equation fitted using factors already incorporated. In
order of importance, Power found the number of plant species on islands was the best
single predictor variable, accounting for 67% of the total variation in bird species
diversity. Island area, on the other hand, accounted for only 33% of variation if
considered alone. It seems logical that habitat heterogeneity (or diversity) for bird
species would be measured by the biotic diversity of their residences, food sources, or
residences of their food resources, i.e. the diversity of the plant community.

Having used plant species diversity as the first factor entered stepwise into the
model, what was the next most important factor? Not area! Once effects on plant
species diversity are removed, area doesn't even account for a significant fraction of
remaining variation, let alone be the most important factor in residual variation. Only one
factor is significant in explaining residual variation of a model including plant species
diversity. That factor is isolation (distance), which accounts for 14% of total variance
when entered as the second factor in the stepwise model equation. The remaining 19%
of variance is unexplained variation, called error variance; no other factor accounts for a
significant portion of variation out of this residual. This is a solid indication that
measures of habitat heterogeneity (diversity) are the critical predictors of island species
diversity.

Power, however, went further. Since plant species diversity, used as a predictor
variable (or factor) is itself a biological variable, he asked what, in turn, explains plant
species diversity. Using the same stepwise regression method, Power found 3 factors
were significant, and they were 3 factors that correspond closely to those which
Diamond found significant in explaining bird species diversity in the New Guinea
satellite islands: 1) area accounts for the largest proportion of plant species diversity,
68%; 2) latitude accounts for 15% of the variance. Latitude effects are detectable only
after effects of area have been removed, especially since the latitudes of the Channel
Islands don't differ by much. This factor apparently measures position of islands relative
to North-South air and water currents off the California coast, rather than a direct impact
of latitude itself. 3) Isolation, or distance from the nearest source pool, with the islands
all at similar distances from mainland, accounted for only 3%, but was significant. The
remaining 14% of variation was unexplained, i.e. error variance. Taking this analysis at
face value, we can see the origin of path coefficients. It suggests that the underlying
factors which explain bird species diversity on the California Channel Islands are
abiotic, the same factors which Diamond found in his studies, particularly area and
factors important in assessing isolation, but that the proximal factors important to the
birds are biological as well. A statistical path then takes the following form:

area ----> plant species diversity

latitude island area ----> bird species

distance isolation diversity

At this point it should be clear that a major group of biogeographers believe that island
area is an excellent, though indirect, indicator of island species diversity.

While we may be accustomed to thinking of islands strictly in 'geological' terms, it is

clear that islands take many forms, including lakes, forest patches in agricultural lands,
or even zebra mussels colonies. The key aspect of islands is that they are favourable
habitat surrounded by inhospitable habitat. Looking at how zebra mussel colonies on
soft sediments in Lake Erie can be 'islands', Bially and MacIsaac (2000) looked at
invertebrate species diversity in relation to island area. A very clear image emerged.
Small islands (10 cm2) host only about 8 species, on average, while midsize islands
(100 cm2) supported ~13, and large islands (70,000 cm 2) supported about 20 taxa. The
invertebrates utilize gaps between mussel shells as habitat, and mussel feces and
pseudofeces as food.

The Basic Model of Island Biogeography

The model is one of a dynamic equilibrium between immigration of new species

onto islands and the extinction of species previously established. There are 2 things to
note immediately: 1) this is a dynamic equilibrium, not a static one. Species continue to
immigrate over an indefinite period, not all are successful in becoming established on
the island. Some that have been resident on the island go extinct. The model predicts
only the equilibrium number of species, will remain 'fixed'. The species list for the island
changes; those changes are called turnover. 2) The model only explicitly applies to the
non-interactive phase of island history. Initially, at least, we will consider only events
and dynamics over an ecological time scale, and one which assumes ecological
interactions on the island occur as a result of random filling of niches, without
adaptations to the presence of interacting species developing there. Evolution is clearly
excluded.

The variables used in the basic model are I s, the immigration rate, which is clearly
indicated by the subscript to be species specific, i.e. to be dependent on the number of
species already present on the island. Here we're not counting noses, but rather the
rate at which new species (those not already present on the island) immigrate. Phrased
explicitly, it is the number of species immigrating per unit time onto an island already
occupied by S species. Also Es, the extinction rate, measured in species lost per unit
time from an island occupied by S species. Finally, we need to know the size of the pool
of species in the source area available to colonize the island.

The immigration rate Is must certainly decrease monotonically (on average) as the
number of species on the island increases, since as S increases there are fewer and
fewer species remaining to immigrate from the pool P of potential immigrants at the
source. If all species were equally likely to immigrate successfully (i.e. had equal
dispersal capabilities), but actual immigrations were chance events, then the
relationship between Is and S would be linear, the probability of a new species
immigrating would be directly proportional to the number of species left to arrive. There
are, however, considerable differences in the dispersal abilities of species in source
areas. Those with the highest dispersal capacities are likely to colonize an island rapidly
(have a higher immigration rate), and later, on average, those with lower dispersal
capacities will follow. They will not only immigrate later, but the rate at which they
immigrate will be lower because they have lower dispersal capacities. The rate at which
species accumulate on islands is, therefore, initially rapid and then slower. Also, among
those species with lower dispersal capacities the successful immigration of any one
species should have less effect on the immigration rates of species remaining in the
source pool (we have not removed a likely immigrant from the pool) than would the
earlier immigration of a highly dispersible species. Therefore, this part of the curve
should be 'flatter'; the rate of immigration should be little affected by the arrival of one of
these poor dispersers. The result is an observed immigration rate curve which is
concave. The actual (or theoretical) curve for any island is dependent on its isolation.
For any source pool, the observed rate, while similar in shape, will be lower for more
distant islands than for closer ones. Immigration rates are graphed from the left hand
edge of figure 1, declining from the y axis with an increasing number of species already
present.

 
Figure - The basic graphical model of equilibrium in the MacArthur-Wilson model. Figure
from Brown and Gibson -Biogeography.

The extinction rate Es should be, from parallel reasoning, a monotonically

increasing function of S. If area, for example, acts only through its effects on population
sizes, and extinctions are the chance result of small population sizes and demographic
stochasticity, then as the number of species increases, the number of species with
small populations and subject to chance extinctions increases in proportion, i.e. the
relationship would be linear. However, if we consider a more realistic biological
scenario, then as the number of species increases, depressant interactions within and
between species (competition, predation, parasitism) are more likely to occur, and
extinctions are more likely as a result. Remember that these are not species that have
evolved adaptations to interactions. Effects are direct and unmoderated. Since any
extinctions resulting from interaction are in addition to those resulting from demographic
stochasticity, the more realistic shape for the extinction curve is concave upwards. The
extinction rate begins at 0 when there are no species on the island, then increases as
species accumulate. At least for purposes of simplicity in looking at the basic
implications of the model, the extinction curve can be thought of as a mirror image of
the immigration curve.

We now have all the information to produce the basic graphical model. That model
predicts that there is some value of S, which is called Ŝ, for which immigration rate and
extinction rate are in balance; there is a dynamic equilibrium. At that diversity on the
island species are immigrating at a rate equal to disappearances due to extinction. The
result is constant change in the species list on the island; that change in names occurs
at a rate called x, the turnover rate. The length of the species list, however, should
remain constant. This is a stable equilibrium since, should something happen, and the
number of species on the island be perturbed, the imbalance between immigration and
extinction rates at the new S would tend to return island diversity toward its equilibrium
value. Below Ŝ additional species accumulate; immigration rate is larger than extinction
rate. Above Ŝ the reverse is true, extinctions exceed immigrations and the number of
species declines to Ŝ.

Tests of the Model

To test the model, an important piece of evidence is a carefully designed

manipulative experiment studying the fauna which colonize 'islands'. One of Wilson's
students, Dan Simberloff, tested the model using islands which consist of mangrove
mangles in the Florida Keys. Simberloff's Ph.D. thesis had consisted of measurements
of the re-colonization of these islands following 'defaunation' (he had encased individual
mangles in giant plastic bags, sprayed them with short acting, low persistence
insecticides, then followed the rates, numbers, and species which immigrated onto them
after exposure). Re-equilibration, i.e. reaching a stable number of species, had occurred
within 3 years of fumigation in his earlier experiments. In a second series of studies
(Simberloff 1976), the manipulations were equally inventive. After the islands had been
censused, and an equilibrium number of species determined for each island (a 'control'
diversity), crews moved in with chain saws, handsaws and hatchets, and each island
was split into 2 or more smaller parts, with water gaps of 1m between. To the insects,
apparently this 1m gap was sufficient to make crossing from one sub-island to another a
jump dispersal. The smaller, sub-islands were then censused repeatedly over a time
interval sufficient to permit re-equilibration to find out how species numbers changed
with island area. Remember, the area censused had been part of a previous island, and
should contain all habitats (plant parts, vertical structures) in the same proportions as
before (i.e. the same habitat heterogeneity, however measured). Alterations were only
quantitative, in the form of area reduction, no unique feature was removed.

The results were clear-cut. Each island reduced in size re-equilibrated at a lower
insect diversity. Considering all the experimental islands in developing a model for the
pattern in reduction, the diversity change fit a log-log relationship (i.e. a power function)
between diversity and area. Thus, Simberloff's data fit the original species-area
relationship. Area was the key determinant. The process of re-equilibration, however,
involved extinction of species from islands supersaturated due to their reduction in size.
We have already encountered the underlying biological cause of those extinctions:
population sizes of 'marginal' species, that is those whose populations were already
small before reduction in area, were decreased to the point where chance extinction
due to demographic stochasticity became likely, and re-colonization unlikely. Such
extinctions are an important component of the equilibrium model of island
biogeography.

Figure - Effect of island fragmentation on insect diversity in mangrove mangles.

Simberloff (1976).

There are few islands that have been studied over long enough periods to test the
hypothesis of equilibrium with turnover, i.e. the occurrence of a stable but dynamic
equilibrium. Among those few are the California Channel Islands. The interpretation of
these data is a source of continuing controversy. That's important, because the crux of
the equilibrium theory is proof (or documentation) of insular turnover at equilibrium. A
paper (Gilbert 1980) found 25 attempts to document turnover at equilibrium, and found
few (basically just mangrove island studies by Simberloff) acceptable without question.
In Simberloff's original defaunation studies, for example, one island supported 7 species
of Hymenoptera prior to fumigation and 8 after equilibrium had been re-established
about one year later. However, only two of these species were present both before and
after fumigation. This sort of experimental study is designed to allow for rapid re-
equilibration.

The Channel Island studies represent an interesting attempt to deal with the
problems of scale (here time) when dealing with most real ecosystems. Recognizing
that there may be difficulties (the initial, historical survey of species presences on the
island used breeding records collected over many years, rather than a single survey at
one time), Diamond's studies of turnover on the Channel islands are still regularly cited
(Diamond 1969).

Initial data reported collections and observations indicating the fauna of individual
islands in 1917. Diamond compared those species lists with a survey he did in 1968.
Over the 51 years between censuses the numbers of species on islands remained
almost perfectly constant, but turnover was as high as 62%, i.e. as much as 62% of the
original list had been replaced by new species. The islands had the following
characteristics:

Island 1917 1968 Extinctions Immigrations %turnover

Los Coronados 11 11 4 4 36

San Nicholas 11 11 6 6 50

San Clemente 28 24 9 5 25

Santa Catalina 30 34 6 10 24

Santa Barbara 10 6 7 3 62

San Miguel 11 15 4 8 46

Santa Cruz 36 37 6 7 17

Anacapa 15 14 5 4 31

 
These data seem initially to fit the equilibrium theory quite well. Numbers remain almost
constant while turnover occurs in a significant number of species. However, the theory
also suggests, as you will soon see, that turnover should be related to island area
(through effects of area on extinction rates) and/or isolation (through effects on
immigration rates. Neither was the case; instead turnover was approximately inversely
proportional to the number of species present. That is not forecast by the model.

Figure - The number of species in censuses of 3 of the California Channel Islands.

Figure - % turnover in species numbers on California Channel Islands. (a) for nine of the
islands. (b) for Anacapa as a function of time between pairs of surveys.

Why should turnover be related to island area or isolation? Consider first 2 islands
at equal distance from the source, but differing in area. Long distance (jump) dispersal
is generally assumed to be a chance event, not directed or goal oriented. In that case,
dispersal probabilities and immigration rates onto the 2 islands should be the same.
Area, however, does affect the extinction rate of colonists. The larger island should
have 1) higher habitat heterogeneity, 2) decreased intensity of interactions due to
reduced niche overlaps resulting from habitat heterogeneity and 3) larger population
sizes making chance extinctions less likely.

These factors should be operative, at least in a relative way, independent of the

number of species present. Therefore, the extinction curves should have similar shape,
but have lower values for the larger island. Putting this comparison on a graph, but
using a linearized version of immigration and extinction curves, we find a larger
equilibrium number of species on the larger island, but also a lower turnover rate on that
island.

To assess the effects of isolation consider 2 islands of equal size, but located at
differing distances from the source. With identical sizes we assume that habitat
heterogeneity, population sizes and interactions on the islands are quantitatively
identical, and thus they have the same extinction rate curves. Immigration rates onto the
more distant island should, however, be lower at any S since the probability of a
successful dispersal decreases (possibly exponentially) with distance. We can go
further, and suggest that the decrease should be most noticeable for species which tend
to be among the first colonists. Later immigrants with lower dispersal capacities have
only a slim chance anyways, and depend on rare, special conditions like storms for
successful immigration. For these species a change in distance should mean less in
shifting immigration rates. Once more we turn these suggestions into a comparison on
the graph. The more distant island has a lower equilibrium number of species, but also
a lower turnover rate at equilibrium than an island closer to the source.

Figure - Multiple immigration and extinction curves indicating effects of differences in

size and isolation on equilibria and turnover rates. Brown and Gibson (1983).

These comparisons can be combined in various interesting and complicated ways.

Rather than document the possibilities, it is probably more valuable to attempt to list the
assumptions and predictions of the basic MacArthur-Wilson model. Some of the ideas in
this list will not be fully examined until later in this section.

Under What Conditions Does the Model Apply?

1) Islands are real isolates (rescue effect, discussed below, not important)

2) Islands have comparable habitat heterogeneity (complexity). There are no gross

environmental changes over the time period of colonization

3) Species counted on islands are residents

4) There is a definable mainland species pool

What Are the Characteristics of the Equilibrium?

1) It is dynamic

2) It is approached asymptotically

3) The process is inherently stochastic

4) The model and the equilibrium are describing processes in ecological time

What Are the Characteristics of Turnover?

 1) The process is not successional

2) Species replacements occur frequently

3) Immigration rates decrease with increasing species numbers. Extinction rates

increase with increasing species numbers

What Influences the Equilibrium Number of Species?

1) Influenced by area through extinction rates

2) Influenced by isolation through immigration rates

3) Varies faster with area on distant islands (see below)

4) Varies faster with isolation on small islands (see below)

With this summary in mind, we return to problems. With regard to Diamond's data, no
combination of size and isolation leads to the prediction that turnover rate is inversely
(or in any other sense) proportional to the number of species on an island.

Since the data are repeatedly cited and classic, it's worth trying to understand why
this anomalous result was reported. There are a number of possible answers, and
arguments in the literature could be described by indicating that 'the fur has definitely
flown'. For one thing, the interval between the censuses was very long. That may have
had significant effect on the measured turnover. If the time interval is long enough it
becomes likely that some of the species which had gone extinct at some time between
the censuses also re-immigrated during that interval (or the converse). In either case
the measured turnover would underestimate actual rates. To attempt to correct for that
possibility, Diamond and his collaborators went back to the Channel Islands annually
during the early 1970's, and also used thorough data gathered for Farnes Island off
Great Britain. The result of differences in the interval between censuses is evident in
Fig.8 (and reported in Diamond and May 1978). The result for the Farne Islands is
parallel. In either case the apparent turnover decreases rapidly as the census interval
increases. To show you why, consider what happened to the meadow pipit on Farnes
between 1946 and 1974 (May and Diamond 1977). The pipit bred for 2 years, went
extinct in the 3rd, then went through 5 more cycles of immigration and extinction over
the remainder of the period. From annual census records that indicates 11 turnover
events in 29 years, where a census after 30 years would have recognized only a single
extinction, as well as a constant diversity of 6 species on the island. The same basic
pattern applies to the Channel Islands. Instead of turnover rates ranging from 17-62%
(or .34-1.24% per year), annual censuses indicate actual turnover rates of 1-10% per
year, and are about an order of magnitude larger than indicated by to 51 year interval
for most islands.

That's not the only corrective surgery which has been suggested for the theory. It
is also evident that monotonic rate functions (particularly the immigration rate curve)
may be overly simplistic. That should be evident by drawing a parallel between
accumulation of species on an island and primary succession. When an island is newly
formed (frequently volcanic) it has no organic content in (and frequently no) mineral soil.
The first plants must be special sorts that have no requirement for nutrients from the soil
(or possibly no requirement for soil at all); instead they are soil formers, leaving behind
their nutrients extracted from the rock (as well as their bodies) to improve conditions for
later arrivals. Krakatoa, East of Java, was not only a B movie, but a real historical event
in the 1880's. What kind of immigration curve described the relationship between
immigration rate and the number of species on Krakatoa after its formation. Depending
on our definition of immigration (does it end with landing on the island, or require initial
growth to be counted) and extinction (does a species have to reproduce at least once
on an island before we consider its loss an extinction?) either immigration rate or
extinction rate curves could be modified. For simplicity we'll include the modifications in
the immigration rate curve. Now it isn't monotonic decreasing; instead it may have an
initial rising phase representing the additional immigration possible with the formation of
soil. That is the naive logic. Reality isn't quite so simple. Over the first 50-60 years since
eruption (1883) the 'curves' of the number of species accumulated over time for various
plant groups seem virtually straight lines; there is no decrease in rate of immigration
over this time. Similar arguments could be advanced, producing a similar curve shape,
with respect to other trophic levels. Equilibrium diversity and turnover rates can be
affected by these modifications.

Figure - Colonization rates for vascular plants on Krakatau measured in terms of total
number of species (labeled 13), immigration and extinction rates (14), and as functions
of the number of species present (15). From Thornton et al. (1993).

Since Diamond's data from the Channel Island studies suggested an appropriate
relationship between turnover and area-isolation effects, but could not quantify them, an
appropriate experimental test became important. Jim Brown and his wife Astrid
attempted such a test (Brown and Brown 1977). They studied the colonization of thistle
plants by assorted insects and spiders by repeated census following defaunation.
Almost everything fit the basic MacArthur-Wilson model, but... The turnover rate should
be inversely related to island distance (isolation) according to the theory. If we look at
just distance effects on the same island (i.e. area), the nearer island should have a
higher turnover rate. That's not what the Browns found. Instead, whether plant 'islands'
were large or small the turnover rates were higher on more distant islands.

Site 1 Site 2

# of plants Mean # species turnover rate # of plants Mean # species turnover rate

large- 16 3.82 0.67 9 5.25 0.29

near

large- 7 3.78 0.78 9 4.44 0.42

far

small- 56 1.89 0.78 21 2.21 0.69

near

small- 3 1.33 1.00 11 0.80 0.91

far

Based upon a 5 day census interval (remember what Diamond's data ended up
showing about the importance of census interval), turnover rates were consistently
higher on the more isolated plants. This reversal of the expected pattern is explained as
resulting from the effect of repeated immigration of species onto near islands. The
original model was psychologically, if not explicitly, concerned with a degree of isolation
which made such repeated immigrations unlikely. Under real conditions repeated
immigration may be likely, particularly on near islands. Addition of a new immigrant
member into a small population reduces the probability of extinction. This repeated
immigration is, therefore, termed the 'rescue effect'. It could, as I've just suggested, be
presented as affecting immigration or extinction, but since the key effect is on extinction
rates, making extinction less likely on near islands, that's where the curves are usually
adjusted for rescue. When the rescue effect occurs, turnover rates will tend to be
directly proportional to distance. The effect may be evident in data sets as divergent as
these studies of plant 'islands' and Diamond's New Guinea satellite island avifaunas.

Figure - How the rescue effect modifies curves of immigration, extinction, and turnover
as a function of distance.

From predictions of rescue effect occurrence we can make some general,

graphical predictions of how distance effects immigration, extinction and turnover. Note
that we are here graphing these rates against distance, not against species numbers.
The immigration rate declines exponentially with distance, as we've previously seen in a
variety of data. The extinction rate was previously considered as depending on island
area, and unrelated to distance; it would have been a straight horizontal line on this
graph before we considered rescue. Now we recognize that the rescue effect bends the
extinction curve down at low distances. When we combine these curves to estimate
turnover as a function of distance, it has an intermediate peak; turnover is highest at
those distances when both immigration and extinction rates are high. Very close to the
source extinction rates are low; at large distances immigration rates are low.

The next correction to the simple model is one which questions the validity of the
species-area relationship. Are projections of area-dependent extinction valid for the total
range of island areas studied? Of course, I wouldn't be suggesting the question unless
something were amiss. There is a problem on very small islands. MacArthur and Wilson
recognized that possibility in presenting the basic theory, and suggested such islands
were unstable, should have very high turnover rates, and probably not have area-
dependent extinction curves. Basically, they thought any area effects would be masked
by the instability of the islands. The figure in the original monograph showed a split
curve for extinction rates, i.e. an unpredictable area effect. The original data used to
construct that graph was drawn from studies of the ecology of the Kapingamarengi atoll
system in the Carolina Islands in Micronesia (Niering 1963). These small outcrops
appeared to show a threshold in the species-area relationship at about 3.5 acres in
area. It was suggested that the instability was not habitat destruction by physical forces,
but instability in the presence of fresh water. Below the threshold area the water table
on the island is saline; fresh water availability depends totally on frequent rains. Above
the threshold area there is a permanent 'lens' of fresh water, and extinction of plant and
animal species dependent on fresh water becomes much less likely.

Figure - Predictions of extinction rate characteristics on very small islands. From

MacArthur and Wilson (1967)

Modeling Effects of Disturbance on the Equilibrium Theory

It is apparent that disturbance can have important consequences for observed

equilibria or the lack thereof. What is difficult is the fact that disturbance has effects on
the survival and/or reproductive success of individuals. A disturbance, unless
extraordinarily massive, does not affect every member of a population. Modeling on an
individual basis has been difficult or impossible until recently. An Italian group (Villa et
al. 1992) attempted to evaluate the effect of regular disturbance at differing intensities
on the equilibrium. The following were their conditions:

1) Island habitats were equally distant from the 'source', but differed in size, from 50
'cells' (each cell was a potential site for an individual) to 1100.

2) There were 64 species. Each had its own mean lifespan, interval between
reproduction, and a range of clutch sizes from minimum to maximum, i.e. a life
history. Each species also has a relative dispersal capacity.

3) A colonization species pool with relative abundances in the pool set.

4) Colonization occurs by randomly allowing individuals to disperse according to a

negative exponential distribution (but distributions are affected by relative dispersal
distance). They are successful if they land on an empty cell. If so their life histories
determine whether the population grows or goes extinct.

5) Disturbances occur periodically. Intensity varied from 0%-75%, where this probability
was applied to each individual, and determined the likelihood of the individual being
killed by the disturbance.

Some results of this simulation seem about as you might have predicted. Some
are 'strange'. The two figures below show you some of the key results. Figure 11 shows
you their eyeball estimates of the conditions which resulted in equilibrium. Over the 120
time intervals their simulation ran, slow-growing organisms never reached equilibrium
on large islands, but did on small ones when there was no disturbance (indicated by 0
on the x-axis). They could not reach equilibrium on any islands at higher levels of
disturbance. Fast growing organisms (on the right) could reach equilibrium on any size
island in the absence of disturbance, and with the larger population size possible on
very large islands, could even reach equilibrium in the face of moderate disturbance
levels (i.e. levels 2 and 3).

Figure - indications of relative equilibrium (thick lines) in 10 simulation runs for islands
with different numbers of cells (the y-axis) and different intensities of disturbance (the x-
axis: 0-no disturbance, 1-10% effect, 2-25% effect, 3-40% effect, 5-75% effect). Part a
is for organisms with 'slow' growth (low clutch size, longer interval between
reproduction, longer lifespan) and part b for 'fast' growing individuals.

Figure - Species-area relationships for a) slow-growing and b) fast-growing organisms

affected by disturbance. The 3 curves are for no disturbance (top curves), 25%
disturbance effect (middle curves) and 75% disturbance (bottom curves).

Evident in the above Figure is the community level effect of disturbance.

Disturbance lowers the overall number of species resident, but if there is sufficient time
for equilibrium to have been reached, life history makes a difference. At low levels of
disturbance, the slow-growing species attain a higher diversity. At high levels of
disturbance, species are not able to remain around long, and a greater diversity can be
achieved by being a good colonizer (a weed, rapid population growth, etc.). Counter-
intuitively, when the actual fitting values for these curves are assessed, the steepness
of the species area curve increases when disturbance is present, even though overall
diversity decreases. What all this tells us is we need to know more about the effects of
disturbance in real communities. The real world is affected by disturbance on a more-or-
less frequent basis, and conservation models based on an equilibrium paradigm need to
be re-considered to incorporate some indication of the effects of disturbance.

References

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soft sediments in Lake erie and facilitate benthic invertebrates. Freshwater Biology
43:85-97.
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immigration on extinction. Ecology 58:445

Crawley, M.J. and J.E. Harral. 2001. Scale dependence in plant biodiversity. Science
291:864-868.

Diamond, J. 1969. Avifaunal equilibria and species turnover on the Channel Islands of
California. Proceeding of the National Academy of Science 64:57.

Diamond, J.M. 1973. Distributional ecology of New Guinea birds. Science 179:759-69.

Diamond, J.M. and E. Mayr. 1976. The species-area relation for birds of the Solomon
archipelago. Proceedings of the National Academy of Science 73:262-266.

Diamond, J. and R.M. May. 1977. Species turnover rates on islands: dependence on
census interval. Science 197:266-270.

Gilbert. 1980. The equilibrium theory of island biogeography: fact or fiction?. Journal of
Biogeography 7:209.

MacArthur, R.H. and E.O. Wilson. 1967. The Theory of Island Biogeography.
Monographs in Population Biology, Princeton Univ.

May, R.M. 1975. Patterns of species abundance and diversity. in M.L. Cody and J.M.
Diamond (eds.) Ecology and Evolution of Communities. Harvard Univ. Press,
Cambridge, MA. pp.81-120.

May, R.M. 1976. Patterns in multi-species communities. Chap.8 in Theoretical Ecology -

Principles and Applications. R.M. May (ed.) Saunders, N.Y., N.Y. p.151-157.

Niering, W.A. 1963. Terrestrial ecology of Kapingamarangi Atoll, Caroline Islands.

Ecological Monographs 33:131-160.

Power, D.M. 1972. Numbers of bird species on the California Channel Islands.
Evolution 26:451-463.

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Thornton, I.W.B., R.A. Zan, and S. van Balen. 1993. Colonization of Rakata (Krakatau
Is.) by non-migrant land birds from 1883 to 1992 and implications for the value of
island equilibrium theory. Journal of Biogeography 20:441-452.

Villa, F., O. Rossi, and F. Sartore. 1992. Understanding the role of chronic
environmental disturbance in the context of island biogeographic theory.
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island biogeography. Evolution 23:171.

Islands are, and always have been, special places. They are the basic stuff of daydreams, a source
of life for shipwrecked souls, valuable pawns in international diplomacy. And they're especially
special for scientists. For Charles Darwin on HMS Beagle, they were part of an expedition that
“seemed to me to throw some light on the origin of species.” For Alfred Russel Wallace, who
searched for answers to the same mystery, “islands possess many advantages for the study of the
laws and phenomena of distribution….[I]t is not too much to say that when we have mastered the
difficulties presented by the peculiarities of island life we shall find it comparatively easy to deal
with the most complex and less clearly defined problems of continental distribution.”

For those scientists whose work takes them into the evolution and distribution of species, and the
reasons behind the distribution, islands became even more important in 1967 with the publication
of a slender monograph written by two respected researchers. Robert H. MacArthur of Princeton,
an expert at both ornithology and mathematics, and Edward O. Wilson of Harvard, the now-
celebrated ecologist who studied the distribution of ants in Melanesia, delivered The Theory of
Island Biogeography to the public (Princeton University Press, 1967). With this book they
sought to explain the relationships between species abundance and the size and isolation of the
areas the species inhabit. As a rough rule of thumb, Wilson later wrote, “the number of species
(birds, reptiles, grasses) approximately doubles with every tenfold increase in area.” A tenfold
decrease in area would halve the number of species.

The authors called their argument an “equilibrium theory” because it maintains that species
number rests on a balance between immigration and extinction, with island isolation also playing
an important part. An earlier paper they had written on the subject had escaped widespread
attention, as had a similar theory advanced in a 1948 doctoral dissertation by Eugene Gordon
Moore. Such theories, freighted with mathematical calculations, may have been simply ahead of
their time. James H. Brown and Mark V. Lomolino, in their comprehensive book Biogeography
(Sinauer Associates, 1998, 2nd ed.), note that “in the late 1940s, biogeography was dominated by
descriptive and taxonomic approaches; this was not a propitious time for injecting mathematical
theory and ecological concepts.”

Whatever the atmosphere of the 1940s and early 1960s, in 1967 the MacArthur–Wilson theory
was a big hit. The term “new paradigm” was heard throughout the hills and valleys of ecological
research. Scholars set about testing, proving, and disproving the thesis. Today, 36 years after the
theory's publication, it remains a fundamental force in the field of biogeography. Scientists are
using the theory and its descendants in attempts to solve some of Earth's more vexing
environmental problems—those brought on by ever-shrinking area for species habitat, as well as
an ever-growing realization that the world has entered its first anthropogenic wave of extinction,
one caused largely by habitat loss. The size and isolation of islands, and of places that act like
islands, have assumed an urgent new importance.

Occasional rediscovery
The importance of islands has had to be rediscovered from time to time. After its initial
acceptance, the theory came in for some heavy criticism. Several researchers had trouble
replicating its findings, and one, biologist Mark Williamson of the University of York, called it
“trivial.” Asked recently about his comment, which appeared in his 1981 book Island
Populations (Oxford University Press), Williamson says he still feels that way. “My feeling then
(and in 1981) was there was lots wrong with MacArthur–Wilson but that a central concept, that
there is immigration and extinction, is true and pretty trivial for most communities.” He adds, “In
essence, I think MacArthur–Wilson misses most of what is interesting about biotic communities
on islands.”

Daniel Simberloff, who worked with Wilson on one of his flagship island experiments,
complained that too much of the theory's application by others was “overzealous.” Michael
Rosenzweig, a former student of MacArthur's and professor of ecology and evolutionary biology
at the University of Arizona, says that after MacArthur's untimely death from cancer in 1972,
“people started picking on it. These were people who don't understand theory, to a large extent.
They began to say things like, ‘Well, it's obvious.’” Rosenzweig hesitates a moment and adds,
“Somebody taught me a long time ago that ‘obvious’ science after the fact is stuff we wish we'd
done ourselves.” Also, some of the criticism stemmed from the fact that “ecologists in many
instances don't like mathematics and don't like having to think about dynamic processes.”

Lomolino and Brown, in Biogeography, further noted that before 1960, the study of island
biogeography was dominated by the notion that species composition was relatively static “unless
modified by long-term evolutionary processes.” Community structure was based on “unique
immigration and extinction events, and species number was determined by the limited number of
niches available on each island…. Either a species had already colonized the island in question,
or it never would. Once it arrived, the species either found adequate resources for survival or
failed to establish a population.” MacArthur and Wilson went far beyond this approach and
brought into the equation the idea of a dynamic equilibrium of the opposing forces of
immigration and extinction. Species composition could and would change over time, but overall
species richness would remain relatively stable.

The authors never meant their theory to be a one-stop solution to species diversity and
distribution, on islands or anywhere else. MacArthur and Wilson wrote, in their 1967 preface,
“We do not seriously believe that the particular formulations advanced in the chapters to follow
will fit for very long the exacting results of future empirical investigation. We hope instead that
they will contribute to the stimulation of new forms of theoretical and empirical studies, which
will lead in turn to a stronger general theory.”

The authors have gotten their wish. Lawrence R. Heaney, writing in a 2000 discussion in Global
Ecology and Biogeography (vol. 9), argues that phylogenetic diversification is part of the
immigration–extinction mix, too. And Brown and Lomolino, participating in the same
discussion, say the theory “has not kept pace with relevant theory and our growing appreciation
for the complexity of nature…. The discipline of biogeography, itself, is in a state of
disequilibrium. We may again be about to witness another paradigm shift, which will see the
replacement of MacArthur and Wilson's theory.” Few scientists today would dismiss MacArthur
and Wilson's work as “trivial” or wrongminded, though. The theory has endured and given
scientists a great deal of meat on which to chew.

The theory and what followed it have greatly sharpened the ways scientists (and nonscientists)
think about insular communities, whether they be on oceanic islands, remote mountaintops, or
prairie potholes. The idea of species equilibrium may have lost some of its initial punch, but
dynamism has grown in strength. Lomolino, an ecologist at the State University of New York's
College of Environmental Science and Forestry, comments, “The major paradigm shift that
occurred really, at least in my mind, wasn't so much that systems were thought to be in
equilibrium but that they were thought to be dynamic.” This appreciation of dynamism has
played a significant part in scientists' (and even some policymakers') thinking about how to deal
with the present crises of loss of habitat and of species diversity.

Brown, a biologist at the University of New Mexico (and recipient of the 2002 MacArthur
Award of the Ecological Society of America), points out that historical events, such as hurricanes
and glaciation, can knock ecosystems out of equilibrium. This is a variable that was not included
in the neat equations of MacArthur and Wilson. Some scientists wonder if the present change in
global climate will be one such historical event.

Not just oceanic islands

The MacArthur–Wilson theory has also enlarged scientific thinking about what, for the purposes
of studying and protecting species richness, constitutes an “island.”

The dynamic-equilibrium way of thinking has been applied to everything from mountaintops to
forest patches to freshwater lakes to caves and desert springs. Wilson says he's seen parasites and
pathogens in human bodies identified as denizens of habitat islands. But if he had it to do all
over, he still would want to cast the theory in terms of oceanic islands. “It turned out that, for the
time, ‘island biogeography’ was heuristic,” he says. “Most other possible expressions would
have been less precise and would not have had the same heuristic effect. ‘Island biogeography’
succeeded in part because ecologists and systematists and, in time, the conservationists, could
clearly conceive of the world as both natural habitat islands and insular islands as discrete and
analyzable systems. And so they were able to get both into original theory of their own and into
very productive field research. I think that the heuristic value of a theory really has as much to do
with its success as its truth.”

John Terborgh, codirector of the Center for Tropical Conservation at Duke University, feels that
research generated by the theory has been quite useful. “The work that Jim Brown did and Jared
Diamond did and I did back in the seventies opened people's eyes to the threats posed by
fragmentation,” he says. “And we were able to show that so-called continental land bridge
islands that are way in excess of equilibrium when they're detached from connections to the
mainland lose species pretty rapidly—and that through some calculations based on that, we're
able to predict very well the number of species left in recently created forest fragments.

“So there was power in the theory to make predictions about entirely different scales of time and
space. And that is a pretty resounding affirmation of the basic robustness of the theory, I would
say.” (Diamond, of the University of California–Los Angeles, also has written extensively about
island biogeography.)

Terborgh is one of many scientists who have produced what Wilson called “new forms of
theoretical and empirical studies” that would help build “a stronger general theory.” The Duke
biologist says, “The influence of MacArthur and Wilson has been profound and pervasive, and
it's had deep impacts on conservation biology, certainly. What is happening now, in the kind of
work we're doing and other people are doing, is to begin to understand the mechanisms that drive
extinction.” Certainly a major such mechanism, maybe the biggest one, is fragmentation of
habitat.

Perhaps the most exhilarating outgrowth of the 1967 theory has been the recognition that
mountain ranges, especially in the American Southwest, could be islands as well. Brown, who
did much early work on what is called the “sky islands” network, says that the MacArthur–
Wilson theory “made an absolutely enormous contribution to the way we think about islands and
island life systems, and even continental systems and certain kinds of conservation problems.”
But it cannot explain everything, including what's needed to conserve the ecological integrity of
the mountainous refugia.

The sky islands, which cover millions of acres of land in Arizona and New Mexico, comprise
some 27 mountain ranges that once enjoyed a measure of connectivity but that have become
isolated from one another since the most recent glacial period. Peter Warshall, of the University
of Arizona and a student of the region, has written that growing stretches of desert grasslands
and scrub have “limited genetic interchange between populations and created environments with
high evolutionary potential. The resulting sky island ecosystems support many perennial streams
in an arid climate, have a high number of endemic species, and harbor most game species as well
as most threatened and endangered species in the Southwest.” The archipelago runs from
Mexico's Sierra Madre to the Rocky Mountains of the United States.

The ecosystems received their eye-catching name when conservationist and author Weldon
Heald called them sky islands in 1967, the same year the MacArthur–Wilson theory was
published. David Hodges, the executive director of Sky Island Alliance, the Tucson-based
organization dedicated to conserving the region's ecosystems, says Heald's clever choice of
words immediately pushed appreciation of the mountains to a higher level.

“People had recognized the fact that there were isolated endemic populations contained in
different areas,” Hodges says. But Heald “really was the one who was responsible for making
that jump and comparing them to something people really hadn't thought about before: the fact
that we do have islands here. They're inland islands, and they're [sky] islands because they're up
above everything else.”

The Sky Island Alliance and the closely associated Wildlands Project are constructing a
conservation framework that differs from oceanic biogeography in that it focuses on
connectivity. “It's not the actual endemic species that are found on top of the mountains,” he
says, “though we certainly have situations where we try to do what we can to ensure that these
species persist. But one of our concerns is the species that historically have traveled freely back
and forth between the different mountain ranges, using the mid- and lower-level elevations. In
most cases they're utilizing habitats that are found on all the different islands. But they need to be
able to move around, just to keep the genetic flow that's existing within the population, and also
[to reach] areas for expanding territories. There are cases in which they hit three or four different
sky islands at different times of the year.” Other species, such as the black bear, move vertically
rather than horizontally, depending on the season and availability of food.

Fortunately for the alliance, much of the sky islands land is publicly controlled. And, Hodges
says, public land agencies have been generally enthusiastic about the alliance's aims.

A major part of the work of the sky islands and wildlands projects is done under the name of
“rewilding,” or restoring the habitat of the Southwest to a condition where species (especially top
carnivores) can move freely once again. In the Wildlands Project's journal, Wild Earth, is the
statement “We live for the day when grizzlies in Chihuahua have an unbroken connection to
grizzlies in Alaska.” Since rewilding would require some accommodation by the region's
growing human population, the idea terrifies and enrages some critics of the project, who
populate the World Wide Web with screeds calling the conservationists communists, Nazis, or
agents of the United Nations. About “rewilding,” Hodges says, “Sometimes you start out with
buzzwords, and from an advertising perspective and a branding perspective, at the time it seemed
like a good idea. But it's not a word that really caught on with people.”

The value of the theory

There's no doubt, however, that “sky islands” and the image the term invokes have caught on,
and the MacArthur– Wilson theory of island biogeography is largely to credit.

The sky islands and their current problems were created by a reduction in habitat area, one
initiated by nature and augmented by urban sprawl across the desert floor. Wilson says this is
where the theory has proved most valuable, although its authors hadn't planned it that way.

“It's interesting,” says Wilson, “that neither one of us thought much about the possible
applications of the theory to conservation in the sixties. It was only in 1972, at a memorial
symposium of Robert MacArthur's death, that E. O. Willis, of the University of Kansas, and I
wrote a paper on the applications of island biogeography for the planning of reserves.” For that
reason alone, the theory should be remembered as a success, he says, “even though the original
theory has been replaced by much more sophisticated modeling and applications in areas
MacArthur and I never conceived. Clearly, it has served as a means of predicting extinction
rates, and in the demographic principles on which it is built, a tool for investigating the causes of
extinction in shrinking habitats…. That's one of the big consequences of the theory of island
biogeography. It doesn't matter if it's left in pristine condition; if you reduce the reserve enough
in size, even just a moderate amount in size, you're going to start losing species automatically.”

The lesson that habitat reduction equals extinction, thinks Wilson, seems largely lost on the
policymakers who decide where and how large conservation areas will be—at least those in the
United States. “My impression has been that at the level of the present White House and
Congress, relatively little attention has been paid,” he says. There's been more use of the theory's
principles by conservation organizations, both local and global, as well as the World Bank and
United Nations. Three good examples of awareness on the global scale, he adds, would be the
1.5-million-hectare Noel Kempff Mercado National Park in Bolivia, and large new reserves in
Gabon and in Suriname, where new protected areas have been linked to existing ones to create a
large corridor.

But many governments, including Wilson's own, have failed. And, says the coauthor of the
equilibrium theory of island biogeography, his fellow scientists haven't done enough to educate
the public and influence policy. Researchers should not fear being branded as advocates, he says.
“They won't lose their credibility. They'll gain the respect of the rest of the community when
they do it right.”

Just as he and MacArthur predicted back in 1967, says Wilson, the theory of island biogeography
has become outdated by subsequent research and discoveries. But that was the point. “It
definitely has influenced a lot of thinking. If you've got a good theory, that's what it does. It
creates a cycle.

“I believe that the most important practical application has been to alert others to the crucial
importance of habitat size, the predictable decline of species diversity, even in pristine habitat
fragments, and the usefulness of studying species extinction as demographic change in an
ecosystem context. This is extremely important, because of the inevitability of losing species if
you just reduce habitat.

“A lot of people still go around with the feeling that, ‘Oh, all you have to do is put aside a little
reserve somewhere and then you turn the rest of it into farmland. The little reserve has most or
all the species there that were in the whole area before it began to be converted. You're okay;
you've got a little pocket of species there. They'll be safe. We'll take good care of them, and we
won't let any of them be killed off.’

“But they will die off, and eventually die off in substantial numbers. Just automatically. It's
common sense, when you think about it: If you cut it down so that you've got only room for, say,
a couple dozen shrubs of a certain species, and a butterfly species that lives there depends for its
life cycle on a hundred of those, then the bush may persist for a while longer, but the butterfly
will go out very soon.”

Appendices
Undergrads and Island Biogeography

What do undergraduate students know about island biogeography and the years of research and
thought it has engendered? The answer depends on their teachers and curricula, of course. Here
are the comments of one professor of geography, Susan Woodward of Radford University in
Radford, Virginia.

Woodward says that many of her students have never had a course in biology, doubt the
prevailing scientific views on evolution, and would encounter problems with the complex
equations and area–species curves that form the backbone of the MacArthur–Wilson theory. The
undergrads “freak out at addition, let alone calculus,” says Woodward. So she relies heavily on
descriptive teaching. “I hope The Song of the Dodo [the widely praised book by David Quammen
(Scribner, 1996)] gives them a sense of the adventure of science and of learning in general. It
seems to.”

Woodward starts the semester with an overview of biogeographic thought “from inventory and
description to various attempts at regionalization (e.g., zoogeographic provinces, floral
kingdoms, life zones) to the ecological and theoretical biogeography of today.” She uses
MacArthur–Wilson's Theory of Island Biogeography “(1) [as] an excellent example of
theoretical biogeography wherein numbers replace actual species; (2) as a hypothesis that
stimulated two decades of research; and (3) as one of the underlying principles still guiding the
design of parks and preserves.” She and a colleague who teaches land use stay away from
mathematical equations, using “just the generalized graphs comparing small islands and large
islands, near and far islands.”

Will all this result in better-designed protected areas for Earth's shrinking wilderness?
“Ultimately,” she says, “the best design of preserves will probably come from a team of social
scientists and natural scientists who take into consideration not only hot spots and corridors, but
economic development and sustainable human land uses within the matrix.”

View largeDownload slide

Edward O. Wilson coauthored, with Robert MacArthur, the groundbreaking book The Theory of
Island Biogeography. The book's publication in 1967 began an era of research and discussion of
species distribution. Wilson is University Research Professor and Honorary Curator in
Entomology at Harvard University. Photograph: Jim Harrison.

View largeDownload slide

Robert H. MacArthur is codeveloper, with Edward O. Wilson, of an influential general theory of

species distribution called the theory of island biogeography. The Princeton professor, an expert
in mathematics and ornithology, died of cancer in 1972, at the age of 42. His coauthor later wrote
that MacArthur's death “deprived ecology of a creative genius.” Photograph courtesy of
Princeton Seeley G. Mudd Library.

View largeDownload slide

The Hawaiian archipelago is a living workshop in island biogeography. Before Europeans

arrived, wrote Robert H. MacArthur and Edward O. Wilson, the Pacific islands were home to
only about 40 species of birds, and for a long time the extinction and immigration rates were in
rough equilibrium. The arrival of human settlers in the 19th century, along with the species they
introduced, shifted the curve dramatically. Now the islands house an estimated 4500 alien
species of all kinds. From left to right, the islands are Niihau, Kauai, Oahu, Molokai, Lanai,
Maui, Kahoolawe, and Hawaii (the “Big Island”). Sunlight illuminates half of the islands. They
are among the archipelago's dozens of large and tiny islands that stretch for 1500 miles across
the mid-Pacific. Photograph courtesy of Jacques Descloitres, MODIS (Moderate Resolution
Imaging Spectroradiometer), National Aeronautics and Space Administration.

View largeDownload slide

A mosaic of lands, many of them protected as wilderness and national parks and wildlife refuges,
forms what conservationists in the American Southwest hope can become a network of
connectivity for wildlife. The promoters think of the area as a group of terrestrial islands, not
altogether unlike oceanic islands, and they call it the “Sky Islands Wildlands Network.” The
proposed conservation system covers about 17.3 million acres, about half of which are
considered “islands.” The system extends from the Mogollon Rim in central Arizona and New
Mexico to the northern Sierra Madre Occidental mountains in Mexico. Map compiled by Cory
Jones, Sky Island Alliance.

View largeDownload slide

This is the model that started the discussion of island biogeography. Robert MacArthur and
Edward O. Wilson presented it in simplified form, along with many others that included more
complex variables, in a 1963 journal article and four years later in their groundbreaking book.
The curves—one representing the immigration rate for new species and the other representing
the rate of extinction—reflect the flow and ebb of species numbers on a single island. The
authors called the point at which the lines cross an equilibrial species number. From Robert H.
MacArthur and Edward O. Wilson, The Theory of Island Biogeography (Princeton University
Press 1967, 1995). Reprinted with permission of Princeton University Press.