Ecography 31: 401
407, 2008

doi: 10.1111/j.2007.0906-7590.05323.x
# 2007 The Authors. Journal compilation # 2007 Ecography
Subject Editor: Jens-Christian Svenning. Accepted 21 December 2007

Evolutionary species area curves as revealed by single-island
endemics: insights for the inter-provincial species area relationship

Kostas A. Triantis, Moisis Mylonas and Robert J. Whittaker
K. A. Triantis (konstantinos.triantis@ouce.ox.ac.uk) and R. J. Whittaker, Biodiversity Research Group, Oxford Univ. Centre for the
Environment, South Parks Road, Oxford, OX1 3QY, UK.
M. Mylonas, Natural History Museum of Crete, Univ. of Crete, Knosos Av., GR-
71409, Irakleio, Crete, Greece and Dept of Biology, Univ. of Crete, Vassilika Vouton, GR-71409 Irakleio, Crete, Greece.

The species
area relationship (SAR) between different biological provinces is one of the most interesting, but least
explored aspects of the well-known species
area pattern. Following the usage that a biological province is a region whose
species have for the most part evolved within it, rather than immigrating from somewhere else, we propose that islands
can be considered equivalent to biological provinces for single island endemic species (SIEs). Hence, analyses of the
relationships between numbers of SIEs and island area can be used as model systems to explore the form of inter-
provincial SARs. We analyzed 13 different data sets derived from 11 sources, using the power (log
log) model. Eleven of
the SIE
area relationships were statistically significant, explaining high proportions of the variance in SIE numbers (R2
0.57
0.95). The z-values (slopes) of the statistically significant SIE
area relationships ranged from 0.47 to 1.13, with a
mean value of 0.80 (SD90.24).
All the island systems in which SIE represent 50% of species exhibited z-values for the SARs of native species higher
than those deemed typical of archipelagic SARs. The SIE
area slopes are quite similar to those previously calculated for
inter-provincial SARs, indicating that, when speciation becomes the dominant process adding to the species richness of
assemblages, high z-values should be anticipated, regardless of the biogeographical scale of the study system.

The species
area relationship (SAR), the increase in species
number with increasing area, is one of the best-documented
general patterns in biogeography and ecology (MacArthur
and Wilson 1967), yet controversy persists about the best
means of describing the relationship and about the precise
form of different types and/or scales of SAR (Connor and
McCoy 1979, Williamson 1988, Rosenzweig 1995, Losos
1996, Lomolino 2000, Scheiner 2003, Triantis et al. 2003,
Whittaker and Fernández-Palacios 2007). As a central goal
of biodiversity science is to understand the mechanisms that
create and maintain diversity patterns (Brown 1995,
Rosenzweig 1995, Whittaker et al. 2001, Whittaker
2004), these controversies remain relevant for pure and
applied ecology.
SARs are typically described using power functions
(Arrhenius 1921) where the exponent (i.e. the slope of
the log-area
log-richness relationship) is commonly referred
to as the z-parameter. The z-parameter has a simple
mathematical interpretation: it is the rate at which species
accumulate with increasing area. However, there is no
consensus on the biological interpretation (i.e. what evolu-
tionary and ecological processes lead to the observed
accumulation) and scale-dependency (e.g. at which spatial
scales particular evolutionary and ecological processes
predominate) of SARs (MacArthur and Wilson 1967,
Connor and McCoy 1979, Lomolino 2000, Gurd 2006,
Whittaker and Fernández-Palacios 2007).
According to Rosenzweig (1995, 2003, 2004), the
‘‘species
area pattern’’ is comprised of three different
species
area relationships, (four, if the point scale, which
depends on sampling effort, is included), whereby processes
operating at different spatial and temporal scales (Schmida
and Wilson 1985, Crawley and Harral 2001) lead to different
z-values (Fig. 1A). In practice, Rosenzweig’s scheme en-
compasses data sets of two different structures. The first form
employs a nested sampling structure and results in species
accumulation curves (SACs) at two scales: these are the point
(i.e. local) scale and the intraprovincial (i.e. regional) scale.
The second form is where the independent variable is the area
of (to varying degrees) geographically discrete and isolated
land masses, and the dependent variable is the number of
native species found within each area: again there are two
scales in his scheme, the archipelagic (i.e. set of islands) and
the inter-provincial (i.e. between regions). Whittaker and
Fernández-Palacios (2007, Box 4.4, p. 94) term these
archipelagic and inter-provincial relationships ‘‘true’’ island
species
area relationships (ISARs) to distinguish them from
the phenomenologically distinct SACs arising from the

401
Figure 1. (A) Three biological scales of species
area curves, as proposed by Rosenzweig (e.g. 2004). Rosenzweig’s species
area pattern
includes four scales, but the point scale is not illustrated in the graphic. The point and intra-provincial scales comprise species
accumulation curves from a nested sampling system, whereas the inter-provincial and archipelagic scales (termed ISARs herein, for island
species
area relationships) are plots of the number of species found in discrete units of space. (B) The inter-provincial ISAR and the SIE

area relationship exhibit similar z-values (slopes) according to results reported herein. Despite the differences in the spatial scale and the
number of species involved, the two systems exhibit analogous trends of increasing species number with area, since in both speciation is
the major process of species addition.

nested sampling designs (see also Gray et al. 2004). It is on
these z-values of true ISARs that we focus in the present
paper.
For real islands, MacArthur and Wilson (1967) report
archipelagic ISAR z values as typically falling within a range
of 0.2 to 0.35, while more recently, Williamson (1988)
reported a much wider range in z, from 0.05 to 1.132. In
his review, Rosenzweig (2004) essentially supports Mac-
Arthur and Wilson, reporting that values typically fall
between 0.25 and 0.45. By contrast, z-values typically
observed for inter-provincial ISARs have a lower margin of
z 0.6 and range upwards, with most lying around 0.8
1.0
(Rosenzweig 2004), and some exceeding 1.0 (Rosenzweig
1995, 2002; Fig. 1A). These observations have very
considerable significance for conservation science as humans
continue to alter the extent and connectivity of habitats
both locally and globally (Rosenzweig 2004, Whittaker
et al. 2005, Whittaker and Fernández-Palacios 2007). It is
thus important to test the robustness and explore the
properties of Rosenzweig’s species
area pattern model.
Archipelagic ISARs have been studied for real and
habitat islands for many taxa in numerous studies over
several decades, and their properties are thus reasonably well
known (Connor and McCoy 1979). By contrast, general-
isations on the form of inter-provincial ISARs derive from
relatively few data sets, of varying degrees of ‘‘fitness for
purpose’’ (see references cited in Rosenzweig 1995,
pp. 264
283, 2002, 2003, Collins et al. 2002). It is thus
this scale and form of SAR that we focus on in the present
work.
Biotic provinces (/regions) can be defined as self-
contained areas that for the target taxon are relatively
independent from an evolutionary perspective, i.e. wherein
most species are endemic to these provinces (Rosenzweig
1995, Losos and Schluter 2000). The concept of a ‘‘self-
contained area’’ is a simplification given that for all biotic
provinces at least some native species can be attributed to
immigration and have not speciated within the province.
Moreover, the question of the criteria used to delimit a
province has no single objective answer. Rosenzweig (1995,
2001) variously suggests thresholds of 60, 80, or even 90%
endemism might be applied (Rosenzweig 1995, p. 279).
The most stringent threshold of ca 90% would ensure that
evolutionary dynamics dominate the rate processes of a
region or island but would also greatly restrict the number
of study systems available for analysis.
Herein, we focus on the form of ISARs for single-island
endemics (SIEs) exploring the idea that for those species
restricted to single islands, the islands in question can be
regarded as provinces (i.e. evolutionarily self-contained
areas sensu Rosenzweig 1995, 2001). Our selection includes
systems for which the percentage of SIEs varies from 15 to
90%. In terms of processes shaping diversity, we can
(following Rosenzweig 1995, 2001) make a strong claim
for each island being equivalent to a distinct biotic province
where a clear majority of species across an archipelago are
SIEs, whereas for those archipelagos featuring endemic
species but where only a minority are SIEs, the equivalence
of the islands with biogeographical provinces/regions is a
more tentative proposal. Using this approach, and by
focusing on these systems, we can study the evolutionary
contribution to ISARs and gain significant insights regard-
ing the patterns arising when speciation is a significant or
the major source of diversity, as happens in the case of
continental biotic provinces/regions. Our first aim is thus to
explore the consistency in form of SIE
area relationships
across taxa and archipelagos of varying proportion of SIEs.
The body of theory discussed above provides no formal
guide to the variation to expect in regard to the z-values for
different taxa, other than that the variation should be rather
limited, given that by definition SIE species treat their
islands as provinces (sensu Rosenzweig 1995), however, we
do expect that systems having the highest proportions of
SIE should have high z values. Our second aim is to use SIE
data to test the robustness of Rosenzweig’s (e.g. 1995,
2004) generalisation that inter-provincial ISARs are sig-
nificantly steeper than archipelagic ISARs, producing z
values typically falling between 0.8
1.0, and 0.25
0.45,
respectively. We do so by comparing z values for SIE with
those for ‘‘all native species’’ for the same taxa.

402
 7.1104
species to be present (S1) are given. n.s.: not significant (p0.05), *: significant (pB0.01), $: slopes of SIE
area relationships and respective ISARs not statistically different (p0.05). + Cases with
Table 1. Source, geographical location, taxon studied, number of islands, range of single-island endemics richness, percentage of the native species SIEs represent, and range of area for the data sets
analyzed in this work. The values of slope (z) and coefficient of determination (R2) for the SIE
area relationships and for the native species
area relationship analyzed, and the area needed for a single

islands with no SIE included for the estimation of the area needed for a single species to be present. For the cases of Anolis lizards (Losos et al. 2006) and the cichlid fishes (Turner et al. 2001), since the

S1 (km2)
Materials and methods

0.065

0.002

0.003
0.03

1.86
0.03

0.59
30.62
We analyzed 13 different data sets derived from 11 sources

0.40 (n.s.)

0.36 (n.s.)

632
206
Native species
(Table 1) and constructed the respective SIE
area relation-
ships. For each data set, the geographical location of the

R2

0.95*
0.58*

0.70*
0.80*
0.54*
0.61*
0.75*
0.85*

0.88*
0.65*

0.78*
archipelago, the taxon studied, the range in number of SIEs,
the percentage of the total species that SIE represent within
each data set and the range of island sizes are presented in
Slope(z)

0.78
1.12
0.38
0.53
0.36
0.46
0.75
0.33
0.37
0.18
0.40
0.44

0.47
Table 1. In the case of Fijian ants (Ward and Wetterer
2006), the data for the species distribution were updated
according to Sarnat (2006). In the data set of Price (2004)

41.09

37.91

57.81
0.05

0.06

1961.5
1.5
for the Hawaiian flora, in addition to the SIE
area
(km2)
S 1

37

38
relationship for all islands, we also present the SIE
area
relationship for the main (south-eastern) islands of the
SIE species

0.38 (n.s.)

0.26 (n.s.)
archipelago, which are a relatively distinct group from the
much older, low-lying northwestern islands. For the cichlids
2

0.88*
0.84*
0.57*
0.65*
0.85*
0.86*

0.62*
0.71*

0.83*
R

of the African lakes (Turner et al. 2001), only lakes with a

surface area 100 km2 were included in the analysis. For
Slope (z)

0.58
0.60
0.47
0.91
1.13
0.48
0.89
0.61
0.54
1.06

0.80
the Anolis lizards of the Great Antillean islands (Losos et al.
2006) and the cichlid fishes of the African lakes (Turner
et al. 2001) the number of SIE is not available, but since the
90
91
Percentage

vast majority of species are regarded as single-island

of SIE, %

90
22
50
82
65
88
57
50
15
54
53

42
endemics ( 90%, Table 1; Losos and Schluter 2000,
Turner et al. 2001, Alphen et al. 2003, Losos et al. 2006),
we consider the resulting slope as a proxy for the respective
SIE
area relationship.
2
1211

1
3193
73
599
7
700

5
179
1
245

11
107
1
198
2
198

1
126

For all data sets we applied the power function model

(range)

2
36
7
57

1
65
SIE

(Arrhenius 1921), taking the logarithm of both sides of the

equation, i.e. LogS  Logcz LogA (eq. 1), where S is
either the number of native species (in a standard ISAR) or
the number of SIE species (in an SIE
area relationship) and
Area km (range)

102
105 007
14
108 722
11 425
114 524

0.7
10 433
116
10 433
2 594
69 484

364
10 458

44
10 338
117
10 458
0.2
10 437
0.8
10 458

278
2 058

A is the area of the respective island (be it a real island or a

278
2 058

lake). For the five cases out of the 13 studied, where islands
2

with no SIE were included in the original data sources, we

omitted them, since their inclusion can strongly affect the
percentage of SIE exceeds 90% we are taking the native and SIE values as equivalent.

estimated values of z and c when using the standard forms

of data transformation used in the literature to accommo-
islands
No of

date zero data points (which is to use log number of

4
8
7

7
6
8
7

7
9
7
14

26

species0.1, 0.5 or 1) and the aims of our analyses require

that we estimate z values that are directly comparable within
Flowering plants
Flowering plants

the others in this study. We used paired t-test to check for

Anolis lizards
Cichlid fishes

Phanerogams

statistical differences between the slopes of SIE
area

Taxon

Land snails
Arthropods
Arthropods
Orthoptera

relationships and the respective ISARs. In addition, in

Orchids
Diptera

order to explore the implications of varying combinations

Plants
Ants

of z and c value, using eq. 1 we estimated the area needed

for a single species to be present for both the SIE and the
Geographical loca-

Great African Lakes

total species richness of each group. For the cases with

Hawaii (large is-

islands with no SIE, we applied a breakpoint regression

Greater Antilles

Canary Islands

Canary Islands

model (Lomolino and Weiser 2001). All statistical analyses

tion

West Indies
Fiji Islands
Caribbean

were carried out using STATISTICA 6.0 (Statsoft, Tulsa,

Hawaii
Hawaii
Hawaii
Hawaii

Hawaii

OK, USA).
lands)
$Ward and Wetterer (2006)

Results
$+ Ackerman et al. (2007)
Izquierdo et al. (2004)

Izquierdo et al. (2004)

The percentage the SIE represent of the total species

Davis et al. (1997)
Turner et al. (2001)

$Peck et al. (1999)

Losos et al. (2006)

richness for each island group considered, ranged from 15

$+ Nishida (2002)
$+ Nishida (2002)
$+ Cowie (1995)

to 90% (0.589SD 0.25) and in 10 cases of the 13

$Price (2004)
$Price (2004)

equaled or exceeded 50%. The slope value (z), the R2 and

the p-values of the regressions, for both SIE and native
Source

species are presented in Table 1. All relationships were

statistically significant, with the exception of those for
+

403
arthropods and plants from the Canary Islands. For these
two cases, if only the western Canary Islands were
considered, with the exclusion of the older and more xeric
eastern islands Lanzarote and Fuerteventura (Whittaker and
Fernández-Palacios 2007, and see Discussion below), the
respective equations became statistically significant: for
Canarian arthropods, z0.82 (R2 0.87, p0.02, n 
5) and for Canarian plants z 0.92 (R2 0.91, p 
0.012, n 5). In all other cases, the R2 values for the
SIE
area relationships were quite high, ranging from 0.57
to 0.95 (mean of 0.76 and SD90.14, n 11). Overall, the
z-values of the statistically significant SIE
area relationships
in Table 1 ranged from 0.47 to 1.13, with a mean value of
0.80 (90.24). If we included the values reported above for
the western Canary Islands, the mean value remained
practically the same 0.81 (90.23).
Considering those systems for which SIE represent
50% of species, the z-values for their ISARs (i.e. analyses
of all native species) ranged from 0.36 to 1.12, with a mean
of 0.55 (90.25, n 10, Table 1). These values overlap
with, but on average were higher than, those deemed typical
of archipelagic ISARs by Rosenzweig (Fig. 1A, Rosenzweig
1995, 2004).
Setting aside the Anolis and cichlid data sets, for which
we lack separate values of native and SIE species, and
including the values for the western Canary Islands, the
z-values for the SIE
area relationships (range 0.48
1.13,
mean 0.7890.23, n 11) were systematically higher than
those for all native species for the same archipelagos (range
0.20
0.75, mean 0.4590.13, n 11).
Considering the slope values of SIE
area relationships
for three different subgroups of the data sets used, based on
the SIE percentage, i.e. 80, 50
80 and B50%, the
resulting mean values are: 80% SIE, z 0.8790.32,
n 4; 50
80% SIE, z 0.7590.24, n 6; and B50%
SIE (including the values for the western Canary Islands),
z 0.8590.06, n 3. Note, that apart from the two
Canarian data sets, the only other data set for which SIE
represented B50% of the native species is the West Indies
orchids (Ackerman et al. 2007), for which the SIE z-value is
0.80 (Table 1). Although we have too few data to be certain
on this point, the data reviewed herein did not suggest any
systematic difference in z values of SIE
area relationships as
a function of the proportion of native species that have SIE
status. Moreover, in eight out of the 10 cases where the
percentage of SIE equaled or exceeded 50% of the total
species richness studied herein, the slope values for the SIE

area relationship were 0.5 and the mean value for all the
10 cases was 0.80 (90.26).
In only one case, Caribbean phanerogams, was there a
significant difference in z values between the ISAR and SIE

area relationship. However, in all cases, except Anolis and
cichlids data and with the inclusion of the western Canary
islands, z-values for the SIE regressions were higher than
those for the ISAR regressions, and collectively, the means
for the z-values of SIE and native species were significantly
different (paired t-test, t 5.43, p B0.001, n 11). The
results remained the same if we excluded the data for the
Canary Islands (paired t-test, t4.67, p0.002, n 9).
404
The extrapolated minimum area threshold for a single
species to be present (S 1), was for all cases higher for the
SIE compared to the total assemblage of native species, with
the greatest differences for Caribbean and Hawaiian floras
and the orchids of West Indies (Table 1).
Discussion
It has previously been noted that ISARs for highly isolated
archipelagos (with a rate of speciation higher than the rate
of arrival of new colonists) can be regarded as being
determined predominantly by evolutionary dynamics,
rather than an immigration/extinction (ecological) equili-
brium (Williamson 1988, Adler 1992, Cowie 1995,
Rosenzweig 1995, Peck et al. 1999, Heaney 2000, Emerson
2002, Cameron et al. 2007), and the expected z-values
should have a lower limit of ca 0.6, reaching unity or even
higher values (Rosenzweig 1995, 2003). Nevertheless,
available data for such archipelagos, where speciation is
the predominant process of species accumulation, are
relatively scarce.
It is clear that the number of SIE cannot be considered as
a direct measure of speciation on islands, rather they should
be thought of as an evolutionary dynamics metric (Whit-
taker et al. 2008). Commonly, species endemic to single
islands can arise through intrinsic factors within the
respective island and/or as a consequence of extrinsic factors
resulting in extinction or speciation events on other islands:
1) some current SIE species may have originated on another
island (or land mass), from which they subsequently became
extinct; 2) some former SIE species may have gone extinct
from the island in question; 3) some former SIE species
may have colonised another island(s) to become a multi-
island endemic (Emerson and Kolm 2005, Witt and
Maliakal-Witt 2007, Whittaker et al. 2007). Moreover,
future inventory or taxonomic work may result in changes
in designation or of known distributions: a general
qualification in all such biogeographical analyses. Never-
theless, the vast majority of single island endemics in the
island systems included in the present analysis may be
considered to have arisen through in situ evolution on
individual islands [(see the respective publication for each
data set and references therein; Table 1; for the Hawaiian
plants see also Price and Wagner (2004) and for the snails
of Hawaii see Holland and Hadfield (2004), Rundell et al.
(2004)]. It is clear that the current analysis would be further
strengthened if we had available molecular phylogenetic
information for all the species considered here (but see
Emerson and Kolm 2007). This would allow us to be more
certain about a SIE arising through in situ speciation and
not through other processes (see above). Similarly, we might
be able to trace dynamics involving species losing their
former status as SIEs by having colonized another island or
islands within the archipelago. Nonetheless, the consistency
of the patterns in z values for the SIE data considered
herein, does provide some confidence that they do indeed
reflect the form of evolutionary dynamics invoked by
Rosenzweig as pertaining to biotic provinces. In addition,
in most of the cases studied (and all but one with a
significant SIE
area relationship), SIEs represent more than
half of the total species present on each island group and in
four cases they represent 80% of the total species richness
(Table 1). Consequently these systems can be considered as
microcosms for evolutionary processes (Emerson 2002).
Our analyses have revealed z-values for all the native
species higher than those deemed typical of archipelagic
ISARs (0.25
0.45; Fig. 1A). Moreover, the z-values for the
SIE
area relationships are also in each case higher than
those for all native species for the same archipelagos.
Consequently, where evolutionary dynamics dominate,
the slope of the ISAR is higher than where ecological
dynamics dominate, although comparisons across different
taxa and archipelagos reveals overlap in z-values. Losos
(1996) and Losos and Schluter (2000), in their work on the
Caribbean Anolis lizards, argue for a spatial threshold where
speciation becomes more important than immigration for
Anolis diversity. At this threshold, the slope of the ISAR
becomes steeper, reaching values above 0.70. They conclude
that, ‘‘the slope of the evolutionary SAR should be higher
than that resulting when immigration is the sole source of
new species’’ (Losos and Schluter 2000, p. 848).
Even though a declining extinction rate with increasing
island area may be sufficient to produce a positive species

area relationship (Connor and McCoy 1979, Ricklefs and
Bermingham 2004, Cadena et al. 2005), Losos and Schluter
(2000) demonstrate that such an explanation is insufficient
to account for very high z-values in their study system.
Estimates derived from fitting the phylogeny of Anolis
lizards to a birth
death model indicated that the extinction
rate was below that for speciation, and far too low to
produce an apparent pattern of increased speciation rate on
large islands. In addition, the results of Losos and Schluter
(2000) indicated that (allopatric) speciation is continuous
and frequent on the largest islands, but rare on the smaller
ones. Similar results are reported for the cichlids of the
African lakes (Seehausen 2006). Consequently, in such
isolated systems, the high z-values of ISARs is the
anticipated result of speciation being the prevailing process
of species addition. Extinction may contribute to positive
ISARs, but cannot alone lead to extreme high z-values.
Accordingly, as larger islands are expected to exhibit higher
speciation rates and lower extinction rates compared to
smaller islands (as Whittaker et al. 2007, 2008), steep slopes
of ISARs should be indicative of systems in which
speciation is the major factor.
The results of the present work confirm the anticipated
relationship of the minimum area thresholds between SIE
and the total species assemblage: the minimum area for SIE
is higher in all cases. The minimum area required for at
least one single-island endemic species to be present cannot
be considered as a direct measure of an island’s size required
to establish within-island diversification, as in some cases,
this involves extrapolation outside the empirical range of
the data: this is dangerous given that the distribution of
values in several cases suggests that a non-linear form
(steeply rising towards an asymptote) might be more
appropriate than the power function. Nevertheless, the
values estimated are an indication of the spatial scales over
which the evolutionary processes are acting, regarding the
taxon and island group under study.
The relatively restricted number of publications studying
the number of SIE in relation to area, and the possibly
confounding effects of other variables such as island age
(Peck et al. 1999, Whittaker et al. 2007, 2008) limits our
ability to detect ecological trends related to the minimum
area required for a SIE to be present. Nonetheless, in our
analysis lower values were found for invertebrates than for
vertebrate taxa, in accord with expectations about their
respective resource requirements and the scale over which
they might form reproductively isolated subpopulations,
leading to new species formation (Table 1). Values for
plants were intermediate, with the exception of the 1961.5
km2 found for the West Indian orchids (Table 1). This is
explicable in relation to the their potential for long-distance
dispersal given that nearly all orchid species have dust-like
and wind-dispersed seeds and that they are known to be
effective colonists of moderately remote islands (Whittaker
et al. 1997).
According to our results, regardless of the SIE percen-
tage, the mean z-value for the SIE
area relationship remains
in all cases steep, and higher than 0.6. Thus, theoretically,
for SIEs arising by in situ speciation on individual islands,
each island might be considered equivalent to a distinct
biotic province, since they originated within these islands.
The provincial dynamics depend rather more on an
evolutionary interplay between speciation, migration and
extinction rates, than on an ecological balance between
immigration and extinction rates (see also Heaney 2007,
Whittaker et al. 2008). The cumulative difference between
the creative process of speciation and the destructive process
of extinction determines the number of species alive in a
biological province and should produce ISARs with high z-
values (Rosenzweig 1995, 2003). The only two data sets
that failed to produce a significant SIE
area relationship
were the Canarian plant and arthropod data, for which the
overall percentage of SIEs were 22 and 15% respectively
(Table 1). Whilst this might indicate that they are not
acting as biotic provinces for these taxa, it could equally
reflect on the environmental characteristics and develop-
mental histories of the archipelago (Whittaker et al. 2007,
2008), as exclusion of the two older, large, low-elevation
and largely arid eastern islands of Lanzarote and Fuerte-
ventura resulted in significant z values of 0.77 (arthropods)
and 0.89 (plants) for the remaining five islands. The
particular data sets indicate that there might be cases where
area by itself will fail in effectively describing the evolu-
tionary dynamics of the system (Whittaker and Fernández-
Palacios 2007, Whittaker et al. 2007, 2008).
Consequently, according to theory, the resulting z-values
of SIE
area relationships should be similar to those arising
when different biogeographic provinces are compared (Fig.
1B). The resulting mean z-value from the present work
(0.8090.24) is almost identical to the mean z-value
(0.7090.23) arising from the 13 inter-provincial ISARs
presented by Collins et al. (2002, Table 1, p. 459).
Including a further set of four z-values of inter-provincial
ISARs presented by Rosenzweig (2002, 2003) and not
considered by Collins et al. (2002), the mean value is 0.79
(90.29), which is even closer to the mean arising from the
SIE
area relationships in the present study.
As noted above, every real province is ‘‘a self contained
region whose species originate entirely by speciation within
405
the region’’ (Rosenzweig 1995). Rosenzweig notes that this
definition sets up ‘‘a continuum between islandness and
province-ness’’ and as the proportion varies for 100%
immigration toward 100% speciation, the region is passing
from island to province. Thus, there is not a clear threshold
for a region to be considered as a province, but percentages
60% endemism and surely close to 80
90% should
be enough to allow speciation
extinction dynamics to
dominate the rate processes of such regions (Rosenzweig
1995, 2001). As already mentioned above, for the 10 cases
where SIE percentage exceeds 50%, the mean z-value is
0.80 and for the four cases where the percentage of SIE
exceeds 80%, the mean value is 0.87. Moreover, for the
three cases in which the percentage of SIE equals or exceeds
90% (Table 1), the mean z value is unity (1.009 0.21).
Hence, the results of the present work provide significant
support for the proposition that z-values from inter-
provincial ISARs should be very high, approaching unity
(Rosenzweig 1995, 2001). ‘‘As provincial area grows,
species diversity increases almost linearly’’ (Rosenzweig
2001, p. 362). This should hold not only for the scale of
recognized global biogeographic regions (as Gaston and
Hudson 1994, Collins et al. 2002) but also for any system
in which speciation is the major process. Of course, the time
scales in which species evolve and go extinct may differ
between island systems and global biogeographic provinces.
Nevertheless, it seems that the two system types exhibit
analogous patterns of species accumulation with area.
The law that the inter-provincial SARs follow is clear
and it appears that diversity in provinces has been following
such a law for hundreds of millions of years (Rosenzweig
2001, 2003). On the other hand, Collins et al. (2002)
pointed out that area does not by itself explain most of the
variation in species diversity among provinces (and see
Hawkins and Porter 2001). There are two main points here.
First, even though our knowledge about the species existing
in our planet has greatly increased during the last decades,
with the description of many new species, we are still a long
way from complete species inventories at the provincial
scale, with the exception of some major taxa (Whittaker et
al. 2005). Hence, inter-provincial ISARs could suffer from
this lack of complete inventories, affecting the variance of
the species richness explained by area. Second, and as
importantly, it is well established that many other factors
(e.g. climate, habitat diversity, evolutionary history, pro-
ductivity) can strongly influence provincial species diversity
(Ricklefs and Schluter 1993, Hawkins and Porter 2001,
Whittaker 2004). And, within oceanic islands, variation in
system properties in relation to time and the life cycle of
islands is also important to a complete understanding of
patterns of diversity and endemism within archipelagos
(Whittaker et al. 2007, 2008). However, even though area
may be an incomplete and incompletely understood
predictor of species richness, so far it has proved to be the
most powerful single explanatory variable for systems across
which area itself varies substantially (Whittaker and
Fernández-Palacios 2007). Using SIE numbers in well-
defined island groups, we keep the variance and influence of
variables other than area as close to constant as practical, so
that the influence of area can be determined and explored.
Area explains an average of 76% of the species richness
variance in the systems studied in the present work.
406
Therefore, studies of evolutionary dynamics in relation
to area, employing data for single island endemics (along-
side the consideration of available phylogenetic informa-
tion), would be well worth pursuing in other taxa and
regions, as they could be used as model systems to test: 1)
variation in critical island sizes below which within-island
diversification does not occur; 2) how these thresholds vary
with taxa and island groups; 3) how consistent the form of
inter-provincial ISARs are; 4) their capability for predicting
future diversity; and they may also be used 5) to recognize
and examine the influence of additional factors such as
climate, productivity, etc. Such studies, can offer great
insights into basic questions of conservation biogeography,
such as the potential impact of habitat fragmentation and
loss and homogenization on biological diversity (Mac-
Arthur and Wilson 1967, Collins et al. 2002, Rosenzweig
2003, Whittaker et al. 2005).
Acknowledgements
We are grateful to our colleagues A.
Parmakelis and S. Sfenthourakis and especially to Gregory Mayer,
Jonathan Losos and Robert Ricklefs for their constructive
comments on earlier drafts of the manuscript. Jens-Christian
Svenning and two anonymous referees made valuable suggestions,
which led to significant improvements in the manuscript. The
present work was supported by the Marie Curie Intra-European
Fellowship Program (project ‘‘SPAR’’, 041095).
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