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RESEARCHPAPER
One size does not fit all: no evidence foran optimal body size on islandsgeb_531 1..10Pasquale Raia1,2*, Francesco Carotenuto1 and Shai Meiri3
1Dipartimento di Scienze della Terra,
Universit Federico II, Largo San Marcellino
10, 80138 Naples, Italy, 2Center for
Evolutionary Ecology, Largo San Leonardo
Murialdo 1, 00146 Roma, Italy, 3Department
of Zoology, Tel Aviv University, Tel Aviv 69978,
Israel
ABSTRACT
Aim Optimal body size theories predict that large clades have a single, optimal,body size that serves as an evolutionary attractor, with the full body size spectrumof a clade resulting from interspecific competition. Because interspecific competi-tion is believed to be reduced on islands, such theories predict that insular animalsshould be closer to the optimal size than mainland animals. We test the resultingprediction that insular clade members should therefore have narrower body sizeranges than their mainland relatives.
Location World-wide.
Methods We used body sizes and a phylogenetic tree of 4004 mammal species,including more than 200 species that went extinct since the last ice age. We tested,in a phylogenetically explicit framework, whether insular taxa converge on anoptimal size and whether insular clades have narrow size ranges.
Results We found no support for any of the predictions of the optimal size theory.No specific size serves as an evolutionary attractor.We did find consistent evidencethat large (> 10 kg) mammals grow smaller on islands. Smaller species, however,show no consistent tendency to either dwarf or grow larger on islands. Size rangesof insular taxa are not narrower than expected by chance given the number ofspecies in their clades, nor are they narrower than the size ranges of their mainlandsister clades despite insular clade members showing strong phylogeneticclustering.
Main conclusions The concept of a single optimal body size is not supported bythe data that were thought most likely to show it.We reject the notion that inclusiveclades evolve towards a body-plan-specific optimum.
KeywordsBody size evolution, Brownian motion model, island rule, mammalianphylogenetic tree, optimal body size theory, phylogenetic dispersion.
*Correspondence: Pasquale Raia, Dipartimentodi Scienze della Terra, Universit Federico II,Largo San Marcellino 10, 80138 Naples, Italy.E-mail: [email protected]
INTRODUCTION
Optimal body size theory (hereafter OST) suggests that large
clades (e.g. mammals) have a fundamental size at which fitness
is maximized (Maiorana, 1990; Brown et al., 1993). In
mammals this size was claimed to be 100 g, based on interspe-
cific allometries of resource acquisition and reproduction
(Brown et al., 1993), although a different optimum (1 kg),
based on a different model, was empirically estimated by
Damuth (1993). The applications of the OST have been
extended to birds and snakes, although an optimal size based
on fitness estimates (33 g) has been calculated only for birds
(Maurer, 1998).
Brown et al. (1993) asserted that the OST explains the global
body size frequency distribution of terrestrial mammals which,
they claimed, shows a strong mode near 100 g. Moreover, they
stated that the OST explains the tendency of small mammals to
grow large and of large mammals to grow small on islands (i.e.
the island rule; Van Valen, 1973). Interspecific competition was
the only force suggested to keep species away from the body size
optimum. Brown et al. (1993) and Damuth (1993) hypothesized
that on islands, where faunas have few species, interspecific
competition is reduced and species are free to evolve towards
their optimal size, driving the island rule.
Although simple and intuitively appealing, the OST has
received as much criticism as support. Optimal size theory was
Global Ecology and Biogeography, (Global Ecol. Biogeogr.) (2010)
2010 Blackwell Publishing Ltd DOI: 10.1111/j.1466-8238.2010.00531.xwww.blackwellpublishing.com/geb 1
claimed to be based on unrealistic parameters (Kozlowski,
1996), to mix reproductive output and conversion rate (Perrin,
1998), individual with population estimates (Kozlowski &
Gawelczyk, 2002) and be inherently inconsistent (Bokma, 2001;
cf. Brown et al., 1996).
Empirically, the OST failed to predict the allometric scaling of
life-history traits in bats (Jones & Purvis, 1997; Purvis, 2006)
and insectivores (Symonds, 1999), the mass distributions of
Australian marsupials (Chown & Gaston, 1997), and the rela-
tionships between body size and home range in strepsirhine
monkeys (Lehman et al., 2007). On conceptual grounds, the
existence of a single optimal size at 100 g implies that all
members of successful mammalian clades such as artiodactyls
and carnivores are suboptimally sized, which is a contentious
assertion (Blackburn & Gaston, 1996; Meiri et al., 2005).
Within the realm of island biogeography, the evidence for
OST is usually estimated from the relationship between body
size range and island area: the body sizes of the smallest species
on each island are regressed against island area (a proxy for
species richness), and so are the body sizes of the largest species
on each island (see, e.g., Marquet & Taper, 1998, Fig. 1; Boback
& Guyer, 2003, Fig. 2; Meiri et al., 2005, Fig. 2). OST predicts
that the intersection of these two regression lines will occur near
the optimal size. Larger islands may contain more extreme sizes
by chance alone, as more species are drawn from the species
pool (Marquet & Taper, 1998). Thus the regression lines will
always intersect at some small island area value, and, by chance,
this area would probably correspond to a body size near the
taxon mode. We therefore argue that it is reasonable to assume
that, under the OST, the true intersection should be nearer to the
optimum than a random draw of species from a global pool
according to the richness on each island.
The intersection of such regression lines was near the putative
100 g optimal size of terrestrial mammals (Marquet & Taper,
1998) and near the modal size of snakes (0.881.08 m; Boback &
Guyer, 2003). Interestingly, however, the intersection of the
regression lines for snakes (1.22 m) was further away from the
modal size than randomized data (0.92 m, Boback & Guyer,
2003). Likewise, for mammals, Marquet & Taper (1998) found
that mass at the regression line intersection (51 g) was signifi-
cantly lower than expected by chance (100112 g), and thus also
lower than the supposed 100 g optimum. For insular carnivores
Meiri et al. (2005) found that the actual regression line intersec-
tion (4363 g) was far from 100 g, and significantly higher (i.e.
further away from 100 g) than the null (3122 g). Thus island
biogeographic data seem to show that animals on species-poor
islands have non-random body sizes, but not necessarily close to
the putative optimal size of a clade.
The OST was further invoked to explain body size patterns in
insular turtles and bats (Lomolino, 2005), but failed to explain
size evolution of large herbivores and carnivores (Raia & Meiri,
2006). Lomolino (2005) did not directly test the OST, but
claimed nonetheless that his data (for reptiles, mammals and
birds) are consistent with the notion that an optimal size drives
the island rule. He argued, however, that multiple optimal body
sizes exist, and suggested optima are body-plan specific.
Because adherents of the OST claim deviations from optimal-
ity are the result of interspecific competition (only) they predict
that insular mammals, especially on small islands, should often
converge on the optimum. The optimal size is therefore an evo-
lutionary attractor. At least two explicit tests of the existence of
an evolutionary body size attractor (although neither used
insular animals) were presented: Alroy (1998) used putative
ancestordescendant relationships of North American Cenozoic
mammals. He found no evidence that the 100 g body size was an
evolutionary attractor. Roy et al. (2000) found remarkably con-
sistent modes and medians in body size frequency distributions
of four assemblages of marine bivalve species along a latitudinal
gradient in the north-eastern Pacific continental shelf. Yet, using
fossil data, they found no evidence that bivalve genera evolved
toward any optimal size.
We contend that the OST predicts that size ranges on islands
will be narrow, because this theory assumesmany insular species
encounter little competition, and are thus free to evolve towards
the optimum. Some theory of community assembly, however,
predicts that much of the size range will be present even at
relatively low richness: Brown & Nicoletto (1991) and Cardillo
(2002) have shown that species-poor assemblages often have
similar size ranges to species-rich ones, with a size frequency
distribution tending towards log-uniformity in the former
versus log-normal or even a right-skewed distribution in the
latter. Species-rich assemblages mostly differ from species-poor
ones because they contain more modal-sized species (Olson
et al., 2009). Thus OST will predict that size ranges will be more
restricted on islands than expected if insular species are a
random sample of the global species pool. If even species-poor
assemblages cover much of the size range, however, insular
species will have wider size ranges than expected under such a
nave null (Meiri & Thomas, 2007).
Body size is often evolutionarily conserved. Thus, if insular
taxa are phylogenetically closely related, then they are likely to
have a more restricted size range than a random group with the
same number of species on the mainland. A relevant null
hypothesis should therefore account for phylogenetic relation-
ships, because the OST predicts that stabilizing selection for the
optimal size does occur. This implies that insular species will be
more similar to each other than expected under the Brownian
motion model of evolution (i.e. they should evolve body size
conservatism and not just show a phylogenetic signal in body
size; Losos, 2008). The Brownian motion model is one, and the
simplest, of a class of models proposed to account for non-
independence of species trait data due to shared ancestry
(Freckleton et al., 2002). In it, the expected phenotypic diver-
gence between species is proportional to the time since their
divergence. More complex models can account for an early
burst (i.e. non-gradual) mode of trait evolution, or test for the
existence of adaptive peaks (Lavin et al., 2008). The evolution of
mammalian body sizes in the class as a whole probably differs
from a Brownianmotion-like body size evolution in being partly
punctuational (Mattila & Bokma, 2008). Furthermore, ecologi-
cal components such as character displacement, species sorting
in local assemblages, or clinal adaptation to environmental con-
P. Raia et al.
Global Ecology and Biogeography, 2010 Blackwell Publishing Ltd2
ditions (e.g. Bergmanns rule) are known to affect the variance
in body size in mammals even after the effect of phylogeny (i.e.
shared ancestry) is accounted for (Diniz-Filho & Bini, 2008;
Diniz-Filho et al., 2009). The Brownianmotionmodel, however,
is an unbiased depiction of size evolution. It is free from
assumptions regarding the direction of natural selection, and
therefore provides a convenient null to test for the occurrence of
selection for an optimal size as predicted by the OST.
Here we explicitly test two predictions of OST: that insular
species evolved toward the 100 g body size optimum and that
body size range is narrower in insular taxa than expected by
chance. Using a relatively complete body size dataset and phy-
logeny for mammals, we test whether the median size of insular
clades is closer to 100 g, as per Brown et al. (1993), or to 1 kg, as
per Damuth (1993), and other possible values (see below) than
the body size of species in their mainland sister clades. We
further test whether the size ranges of the insular clades are
narrower than those of their mainland sister clades controlling
for clade richness.We compare size ranges in insular clades with
those produced by simulations performed under the Brownian
motionmodel of evolution. Finally, we estimate whether the size
range computed over all insular endemic species within a family
is narrower than expected by chance, after controlling for species
richness.
MATERIALS AND METHODS
Phylogenetic and body size data
To test the predictions of OST using phylogenetically informed
null models, we modified the mammal species-level phyloge-
netic tree published by Bininda-Emonds et al. (2008). We
omitted marine mammals (cetaceans, sirenians and pinnipeds)
and species for which we had no body size data. The modified
tree includes 4004 extinct and extant mammal species. Body
masses were taken from Smith et al. (2003) and additional
sources (see Appendix S1A in Supporting Information). Insu-
larity data are from Smith et al. (2003), verified using multiple
sources. Body size (in grams) was log 10 transformed in all
analyses.
Human activity drove many insular mammals to extinction
(e.g. Diamond, 1982). Alcover et al. (1998) estimated that 27%
of insular endemic mammals have gone extinct since human
colonization of their islands. Human-driven extinctions on the
continents were no less severe. Worse, extinctions were strongly
size-biased (i.e. most extinctions were of large mammals; e.g.
Johnson, 2009). Thus, comparing mammal sizes on islands and
themainlandmay be biased by poor and non-random sampling.
Consequently, we included all species known to have gone
extinct since the final part of the last glacial phase (some
40,000 yr bp), for which a consistent body mass estimate and a
clear phylogenetic hypothesis were both available. We thus
restricted ourselves to species that were still living when human
activities began to affect mammalian faunas world-wide
(Johnson, 2009). These extinct species include disparate taxa
such as marsupials, ungulates, xenarthrans, bats and rodents
(Appendix S1A). Thus, it is reasonable to assume that our
sample of extinct taxa is unbiased in relation to body size or
taxonomy. Body masses for extinct taxa, their status as insular
endemics and the works used to reconstruct their phylogeny and
body sizes are reported in Appendix S1.
To produce our phylogenetic tree, we used the branch
lengths reported in Bininda-Emonds et al. (2008). For extinct
taxa we used estimated age of separation (split age) between
fossil taxa as reported in the source papers if available (Appen-
dix S1). Otherwise, we assessed the split age between sister taxa
by using their oldest fossil occurrence reported in The Paleo-
biology Database (http://paleodb.org/cgi-bin/bridge.pl). Where
no precise temporal reference was available we placed the split
age at the mid distance between the parental and the daughter
node ages. This procedure minimizes variance in branch length
(Webb et al., 2008). Reconstructed node ages were calculated
with the bladj algorithm in Phylocom (Webb et al., 2008). Our
modified tree includes 4004 species, 746 of which are insular
endemics and 217 of which are extinct (Appendix S1). In most
analyses we used all resolved nodes subtending a sister-clade
pair, where one clade includes only insular endemics and its
sister clade includes only mainland species. We contrasted
body size ranges (defined as the difference between the log-
transformed masses of the largest and smallest species in a
clade) of clade pairs, effectively restricting our analyses to sister
taxa. There are 172 such clade pairs, containing in total 1143
species (Appendices S2 & S3). Where the tree contained both
insular and mainland species in polytomies under a single
node, we arbitrarily defined the insular and mainland species
as sister clades, and the parental node as a clade pair. We then
repeated the analyses excluding the polytomous clade pairs
(see below).
Tests for evolution toward the optimal size
We used the Fisher exact test to compare the frequency in which
the median mass within the insular clade is closer to the 100 g
optimum than that of their mainland sister clade, with a null
expected probability of a 1 : 1 ratio. We repeated this test using
only fully resolved clades.
We further tested for the existence of other possible size
optima serving as evolutionary attractors by comparing the fre-
quency of insular dwarfism and gigantism in mammals of dif-
ferent sizes. We tested whether the median mass of species in
insular clades is greater or smaller than the median masses of
species in their sister clades for mainland median masses of
< 10 g, 10100 g, 1001000 g, 110 kg and > 10 kg. This test also
allowed us to infer whether there is a stronger support for an
optimum at 100 g (Brown et al., 1993) or at 1 kg (Damuth,
1993), because, where the masses in the mainland sister clades
are 1001000 g, the former predicts insular dwarfism whereas
the latter predicts gigantism. If, however, there are clade-specific
tendencies towards dwarfism or gigantism but no optimal size
(Foster, 1964; Meiri et al., 2008), we would expect dwarfism and
gigantism to be independent of any optimum.
No optimal body size in mammals
Global Ecology and Biogeography, 2010 Blackwell Publishing Ltd 3
Tests for decreased size range in insular clades
We defined body size range as the difference in log body mass
between the largest and the smallest species within each clade
(equal to the largest size ratio in the clade). Under the OST the
insular clade is predicted to have a narrower size range than its
mainland sister clade.We used three tests of this prediction. First
we performed an analysis of covariance (ANCOVA) on the body
size ranges of sister clades, with the number of species in a clade
as a covariate and insularity (i.e. whether a clade is insular or
not) as the main effect.
In a second test we simulated body size evolution 1000 times
across the whole phylogeny by using the function evolve.phylo in
the R library ape (Paradis et al., 2004). From each simulation we
drew a body size for each species, computed size ranges of main-
land and insular species and calculated the size range difference
as insular minus mainland size range. We standardized this dif-
ference by dividing it by the simulated mean body size of the
mainland clade.This is an indexof difference in relative size range
between insular and mainland sister clades. The same index was
then computed with real data, and compared with the 95%
confidence limits computed across all the simulations.
Finally, we examined insular size ranges which did not involve
clade pairs, while still accounting for taxonomy: we asked
whether endemics have narrower size ranges than a random
draw of a similar number of species from their family, using only
families with at least 10 insular endemics.We produced 999 null
samples by picking species at random, without replacement,
from the family the insular endemics belong to. We calculated
body size ranges for each randomization and assessed significant
deviations using two-tailed tests.
Body size is phylogenetically conserved within mammalian
clades (Freckleton et al., 2002; Blomberg et al., 2003): closely
related species tend to have similar sizes. If insular endemics are
more closely related than expected by chance (i.e. there is phy-
logenetic clustering) testing for size range in these species would
produce high type I error, since insular endemics would be
expected to be of a similar size whether the OST applies or not.
We tested for phylogenetic clustering using two different metrics
(Webb, 2000; Webb et al., 2002): the net relatedness index
(NRI), which is a standardized measure of the mean phyloge-
netic distance (in terms of sum of branch lengths) between a
given species and all the other species in the sample averaged
over all species in the sample, and the nearest taxon index (NTI),
which is a standardized measure of the mean phylogenetic dis-
tance between any given species and its sister species, averaged
over all species in the sample. Here the sample is all insular
endemics within a family. Significant NRI and NTI values were
estimated via comparison with random distributions produced
by drawing n species from the family pool, where n is the
number of insular endemics in that family. A total of 999 null
samples were produced for significance testing. Other metrics
testing for phylogenetic dispersion within samples of species are
available, but tend to be highly correlated with each other
(Vamosi et al., 2009). Tests for phylogenetic clustering were per-
formed in Phylocom (Webb et al., 2008).
RESULTS
Tests for evolution toward the optimal size
In 95 out of 170 cases (55.8%, two pairs with identical island and
mainland median masses were excluded) the median body size
within the insular clade is closer to 100 g (P = 0.328, two-tailedtest). Although many of the polytomies in our dataset are prob-
ably hard polytomies (originating from multiple island coloni-
zation by a single parental mainland species), we repeated the
test with only fully resolved clades. In this test the median size of
the insular clade is closer to 100 g in 42 out of 82 cases (51.2%,
P = 0.999). Hence, the 100 g body size does not act as an evolu-tionary attractor for body size of insular taxa.
The frequencies of insular dwarfs and giants were similar in
all size classes except for the largest one (Table 1). In the largest
size class, the median body size of insular species is lower than
the median size in their mainland sister clades in all but one
clade pair (the mysterious Falkland Island wolf, Dusicyon aus-
tralis is larger than its extinct mainland congener Dusicyon a-
vus). Thus there is a significant (P < 0.001) tendency for large
(> 10 kg) species to dwarf on islands but to no net trend towards
gigantism in small species, and no identified optimal size.
Tests for decreased size range in insular clades
Forty-three clade pairs include more than one species per clade
(Table 2). The insular one has a narrower size range in 22. The
size ranges of the insular clades is not narrower than that of their
mainland sister clades (Wilcoxon paired test, z = - 0.330, P =0.742). An ANCOVA indicates that size ranges differ between
clades (whole model F2,83 = 24.7, P < 0.001, R2 = 0.373) but thisis due to species richness (t = 7.02, P < 0.001). The effect ofinsularity is insignificant (t = 0.60, P = 0.55). Similar results wereobtained when we used the clade pair that each size range refers
to as a factor (species richness: F = 93.98, P < 0.0001; clade: F =2.81, P = 0.0006; insularity: F = 0.68, P = 0.41). We furtherregressed the mainland minus island size-range difference
Table 1 The number of clades in which the median body size inthe insular clade is larger (insular gigantism) or smaller (insulardwarfism) than the corresponding value in the mainland sisterclade, for mammals of different sizes. Body size classes wereassigned according to the median body size in the mainlandclades. Probabilities are for deviations from equality in a binomialtest. Data are from Appendix S2 in Supporting Information.
Body
size class
Median size smaller
on islands
Median size larger
on islands
Binomial
probability
< 10 g 10 13 0.68
10100 g 31 34 0.80
100 g1 kg 19 15 0.61
110 kg 14 13 1
> 10 kg 19 1 < 0.0001
P. Raia et al.
Global Ecology and Biogeography, 2010 Blackwell Publishing Ltd4
against the mainland minus island species-richness difference.
We contend that a positive intercept indicates that insular size
ranges are narrower unless the island clade has more species
than the mainland one. This, however, was not the case: the
intercept is near zero [0.073, 95% confidence interval (CI):
-0.056 to +0.201, slope = -0.28, P < 0.001]. Hence, our testsdemonstrate that body size ranges within clades are a function
of clade richness, but not of whether or not it is insular.
Insular clades within clade pairs do not show narrower size
ranges than expected in Brownian motion model simulations.
Although size ranges of insular clades were narrower than
Brownian motion predictions in 14 out of 43 clades (Table 3), in
15 cases the size ranges of the insular clade were wider than
expected, again with no apparent phylogenetic bias, for they
include bats, rodents, primates, afrosoricids, xenarthrans,
dasyurids and diprotodonts.
Table 2 Summary distribution of theoccurrence of smaller size range andnumber of species for clades within apair. The total number of occurrencesper category is reported at the bottom.Only clade pairs including at least twospecies per clade are included. Nodescorrespond to those in Appendix S2 inSupporting Information.
Clade pair Smaller size range on No. of species larger on
Apodemus argenteus group Mainland Mainland
Antechinus Mainland Mainland
Axis Island Same
Boromys/Clyomys Mainland Island
Cervus Island Mainland
Chimarrogale Island Mainland
Choloepodini Mainland Island
Crocidura Island Mainland
Galidiinae Island Mainland
Hylopetes2 Island Mainland
Hystrix Mainland Same
Kerivoula argentata group Island Mainland
Lemurs/galagos Mainland Island
Lutra lutra group Mainland Island
Macaca 1 Mainland Island
Maxomys Mainland Island
Melogale Island Same
Melomys Mainland Island
Microtus arvalis group Island Same
Monophyllus/Glossophaga Mainland Mainland
Mus 1 Mainland Same
Mus 2 Island Mainland
Mydaus Island Same
Naemorhedus Island Mainland
Natalus Mainland Island
Neotoma albigula group Mainland Mainland
Niviventer Island Mainland
Peromyscus Island Mainland
Petinomys Mainland Same
Phaulomys Island Mainland
Presbytis Same Island
Rattus rattus group Mainland Same
Rhinolophus Island Mainland
Rousettus Mainland Island
Sorex hydrodromus group Island Mainland
Stenodermatinae Island Same
Suncus Island Mainland
Sundasciurus Island Island
Sus Island Same
Tenrecidae Mainland Island
Trichosurini Mainland Mainland
Tupaia Mainland Island
Uromys Island Island
Totals Smaller body size range Lower number of species
Islands 22 14
Mainland 20 19
Same 1 10
No optimal body size in mammals
Global Ecology and Biogeography, 2010 Blackwell Publishing Ltd 5
Using randomizations, the body size range of insular
endemic clades species is not narrower than expected by
chance in any of the 16 families with at least 10 species of
insular endemics (Table 4). In one family (Tupaiidae) the
insular endemics have a wider body size range than expected
by chance. These results strongly argue against the prediction
of OST that size ranges in insular taxa are narrow. This is espe-
cially striking because insular endemics are phylogenetically
clustered (i.e. they are more closely related than expected by
chance given the family they belong to, Table 4). Among the 16
families, 11 show phylogenetic clustering (two-tailed tests).
Insularity is therefore clearly phylogenetically conservative
Table 3 Comparison of actual size ranges differences within a clade pair with data simulated under a Brownian motion (BM) model ofevolution.
Clade pair
Insular clade
size range
Mainland clade
size range
Mean body size of the
mainland clade
Standardized size
difference
Confidence intervals for
standardized size ranges
according to BM model
Apodemus argenteus group 0.340 0.311 1.502 0.019 -0.095 to 0.073Antechinus 0.830 0.440 1.706 0.229 -0.089 to 0.056*Axis 0.130 0.310 4.695 -0.038 -0.065 to 0.066Boromys/Clyomys 1.050 0.820 1.890 0.122 -0.022 to 0.099*Cervus 0.433 0.620 5.060 -0.037 -0.098 to 0.042Chimarrogale 0.019 0.328 1.574 -0.196 -0.127 to 0.091Choloepodini 0.455 0.070 3.745 0.103 -0.05 to 0.071*Crocidura 1.213 1.630 1.034 -0.403 -0.105 to 0.034Galidiinae 0.410 1.420 3.163 -0.319 -0.189 to 0.007Hylopetes2 0.760 0.780 2.443 -0.008 -0.095 to 0.043Hystrix 0.920 0.400 4.135 0.126 -0.086 to 0.087*Kerivoula argentata group 0.060 0.694 0.662 -0.958 -0.123 to 0.013Lemurs/galagos 3.670 1.390 2.490 0.916 -0.049 to 0.226*Lutra lutra group 0.499 0.300 3.890 0.051 -0.015 to 0.066Macaca 1 0.540 0.130 3.745 0.109 -0.009 to 0.088*Maxomys 0.555 0.430 2.045 0.061 -0.065 to 0.147Melogale 0.001 0.260 3.100 -0.084 -0.073 to 0.081Melomys 0.650 0.160 1.895 0.259 -0.036 to 0.142*Microtus arvalis group 0.038 0.100 1.500 -0.041 -0.052 to 0.05Monophyllus/Glossophaga 0.230 0.170 0.993 0.060 -0.089 to 0.032*Mus 1 0.148 0.030 1.375 0.086 -0.082 to 0.085*Mus 2 0.085 0.380 1.169 -0.252 -0.999 to 0.926Mydaus 0.170 0.310 3.955 -0.035 -0.055 to 0.055Naemorhedus 0.080 0.510 4.565 -0.094 -0.069 to 0.016Natalus 0.440 0.020 0.750 0.560 -0.063 to 0.126*Neotoma albigula group 0.310 0.210 2.293 0.044 -0.101 to 0.033*Niviventer 0.190 0.438 1.925 -0.129 -0.182 to 0.032Peromyscus 0.003 0.740 1.516 -0.486 -0.198 to 0.027Petinomys 1.350 1.270 2.055 0.039 -0.073 to 0.076Phaulomys 0.060 0.440 1.475 -0.258 -0.137 to 0.017Presbytis 0.050 0.050 3.825 0.000 -0.001 to 0.095Rattus rattus group 0.558 0.392 2.145 0.077 -0.058 to 0.053*Rhinolophus 0.380 0.860 1.027 -0.467 -0.078 to 0.01Rousettus 0.320 0.090 1.945 0.118 -0.021 to 0.039*Sorex hydrodromus group 0.000 0.830 0.705 -1.177 -0.178 to 0.002Stenodermatinae 0.270 0.320 1.205 -0.041 -0.073 to 0.058Suncus 0.934 1.764 0.880 -0.943 -0.123 to 0.043Sundasciurus 0.690 0.820 2.177 -0.060 -0.049 to 0.16Sus 0.400 1.180 4.650 -0.168 -0.079 to 0.082Tenrecidae 1.940 0.980 2.217 0.433 0.027 to 0.344*
Trichosurini 1.301 0.700 3.417 0.176 -0.121 to 0.038*Tupaia 0.670 0.450 2.117 0.104 -0.066 to 0.194Uromys 0.380 0.520 2.550 -0.055 -0.046 to 0.099
*Wider than expected.Narrower than expected.
P. Raia et al.
Global Ecology and Biogeography, 2010 Blackwell Publishing Ltd6
within mammalian families, but size ranges are not
narrow.
DISCUSSION
We find no evidence to support the notion of a single optimal
size for mammals. Sizes of members of insular clades were not
closer to the putative optima than those of members of their
mainland sister clades. Nor was there any evidence to suggest
that body size ranges within insular clades are narrower than size
ranges in their mainland sister clades, or compared to the pre-
dicted differences in size range drawn from simulations per-
formed under the Brownian motion model of evolution.
Previous studies failed to find evolutionary size attractors
despite assuming that competitive size displacement was not a
factor in the evolution of the lineages under scrutiny (Alroy,
1998; Roy et al., 2000). Many insular faunas are species poor;
hence islands are often viewed as the ideal places for evolution
toward the optimal size (Maiorana, 1990; Brown et al., 1993;
Damuth, 1993; Lomolino, 2005). Brown et al. (1993) explicitly
stated that OST explains the island rule, since insular species can
evolve toward the optimal size (100 g) given the reduced com-
petition regime they encounter. Here we found no support for
the prediction that body sizes of insular taxa are closer to 100 g
than are the sizes of allied mainland taxa. We identified strong
evidence for dwarfism in large insular mammals, as predicted by
the island rule, but our results indicate that this dwarfism only
occurs in very large (> 10 kg) mammals, whereas OST predicts
dwarfism would start at a much smaller size. Furthermore, we
found no evidence for a net trend toward gigantism in small
mammals, which is predicted under both the island rule and
OST. Dwarfism above 10 kg, however, seems to be a general
mammalian pattern, as the 19 clades showing dwarfism belong
to seven orders spanning the entire mammalian phylogeny
(diprotodont marsupials, xenarthrans, proboscideans, rodents,
carnivores, primates and artiodactyls although for carnivores
we have one case of dwarfism and one of gigantism).
Because it posits that small animals evolve larger sizes on
islands whereas large animals dwarf, the island rule will, inevi-
tably, be associated with some intermediate size at which neither
dwarfism nor gigantism is predicted. Our finding, that large
mammals dwarf, but small mammals do not, as a rule, grow
larger on islands, means that we identify a large size range (all
mammals under 10 kg, some 90% of all species in our dataset)
where no size evolution is predicted, rather than a unique size
value (e.g. 100 g). Thus only very large sizes may be considered
suboptimal on islands, and a four orders of magnitude size
range in which animals will show neither dwarfism nor gigan-
tism is at odds with both the island rule and with optimal size
theory.
Body size ranges are not narrower than expected on islands,
and this result persists when we compare actual clade size
ranges, compare real size ranges within clades with simulated
data, or use the entire phylogeny randomizing the status of
insular endemics within families. Given that we find that insular
endemics are more phylogenetically closely related to one
another than expected by chance, the randomizations within
families are very liberal, but we could still not reject the null
hypothesis. This may hint that insular clades may have wider
distributions than expected, suggesting that insular clades may
Table 4 Patterns of phylogenetic dispersion in the occurrence of taxa on islands and their size ranges within mammal families with at least10 insular endemic taxa.
Family
No. of insular
species NRI NTI
Phylogenetic
pattern Size range
Randomized size
range ( SD)
Pattern in body
size range
P (body size range is
random)
Cercopithecidae 17 7.04*** 3.16** Clustering 0.61 0.78 0.19 Random 0.30
Cervidae 10 2.40* -0.07 Clustering 4.90 1.96 1.72 Random 0.38Dasyuridae 11 3.55** 1.88* Clustering 1.63 2.44 0.84 Random 0.41
Macropodidae 15 2.66* 2.31* Clustering 1.61 3.37 1.64 Random 0.47
Megalonychidae 10 2.43* 2.53*** Clustering 1.40 1.83 0.28 Random 0.14
Muridae 217 16.97*** 4.23*** Clustering 2.29 2.43 0.16 Random 0.37
Mustelidae 11 3.08* 2.32* Clustering 1.59 2.16 0.84 Random 0.32
Phalangeridae 11 1.73 0.18 Random 1.30 2.4 1.45 Random 0.70
Phyllostomidae 13 2.70* 4.35*** Clustering 0.76 1.01 0.32 Random 0.37
Pteropodidae 93 3.52** 0.35 Clustering 1.85 1.83 0.05 Random 0.98
Rhinolophidae 18 -1.26 -1.32 Random 0.94 1.00 0.22 Random 0.80Sciuridae 43 5.92*** -2.88** Clustering 2.03 2.27 0.26 Random 0.38Soricidae 33 0.97 1.64* Random 1.31 1.33 0.25 Random 0.87
Talpidae 10 2.31* 2.47** Clustering 2.11 1.35 0.60 Random 0.36
Tupaiidae 11 0.73 -1.07 Random 2.45 1.52 0.85 Larger than expected < 0.05Vespertilionidae 17 -0.16 0.12 Random 0.92 1.03 0.26 Random 0.93
*Marginally significant, 0.1 > P > 0.05.**P < 0.05.***P < 0.01.NRI, net relatedness index; NTI, nearest taxon index.
No optimal body size in mammals
Global Ecology and Biogeography, 2010 Blackwell Publishing Ltd 7
fill the entire morphological space despite often being less
species rich, in line with the observation that wide size ranges
can form even when species richness is low (Brown & Nicoletto,
1991; Cardillo, 2002; Meiri & Thomas, 2007). More tests are
needed to resolve this issue. We contend, however, that the eco-
logical attributes and particular environment of different insular
species probably play a large role in determining body size evo-
lution above and beyond phylogenetic effects (Diniz-filho et al.,
2009) and possible selection for any optima (Brown et al., 1993;
Lomolino, 2005).
For species originating through insular radiations, a conflict-
ing scenario may be envisioned. An adaptive radiation produces
a large number of new species, which may enhance the intensity
of interspecific competition, enhancing body size differences
between species through character displacement, creating an
overall wider body size range (Schluter, 2000; Davies et al.,
2007). Thus with an in situ radiation, insular clades may be
predicted to have larger ranges than expected by chance. Of the
insular clades we analysed (Tables 1 & 2), insular radiations can
be invoked for Madagascars tenrecs and lemurs, and perhaps
for Indonesian mosaic-tailed rats (genus Melomys), Maxomys
mice, Hystrix porcupines and bare-backed fruit bats (genus
Dobsonia); for New Guinean dasyures (genera Antechinus and
Murexia); and for Antillean cave rats (genera Boromys, Botromys
andHeteropsomys). The vast majority of insular clades, however,
do not represent such radiations, and even the Indonesian and
Antilles examples we cite above correspond to insular endemics
inhabiting different islands within an archipelago, that probably
evolved in isolation. Thus, there is no convincing evidence that
in situ speciation and ensuing competition on islands may have
driven species away from the optimal body size.
Our results are at odds with the idea of multiple, large-clade-
specific body size optima on islands (Lomolino, 2005). We
found no support for the prediction that insular faunas evolve
smaller size ranges on islands, a prediction that is independent
of the optimal value per se. Lomolino (2005) referred mainly to
mammalian orders when suggesting multiple, body-plan-
specific optima and thus our family-level analyses should have
been able to detect them. Thus, we doubt the existence of size
optima in general, whatever these optimal values actually are, or
to which phylogenetic or taxonomic level they are thought to
relate.
We argue that the island rule, whether it is true or an epiphe-
nomenon of the tendency of some clades to evolve either large
or small body sizes on islands (Lawlor, 1982; Meiri et al., 2008),
is better studied by considering contingent factors such as
species biology and the ecological characteristics of island
faunas (Lawlor, 1982; Raia et al., 2003; Raia & Meiri, 2006).
These cannot be captured by one set of allometric equations,
resulting in a one size fits all theory such as the OST. We find
that OST is an unlikely explanation for the evolution of body
size on islands. Similarly, we found no support for the idea that
there is an evolutionary attractor for mammal species evolving
on islands.
The theoretical basis of OST is outside the scope of our dis-
cussion, yet it is remarkable that this theory fails to apply
under the circumstances which best match its predictions (on
islands). We conclude that, while dwarfism in large mammals
is a real net trend, gigantism in small mammals is more idio-
syncratic. The evolution of clade-specific size ranges is inde-
pendent of insularity.
ACKNOWLEDGEMENTS
We thank Ally Phillimore for insightful discussion leading to the
start of this project. We thank Joaquin Hortal and Mark
Lomolino for valuable discussion. Anna Loy, David Currie, Jos
AlexandreDiniz-Filho and three anonymous referees kindly pro-
vided important comments on this manuscript. Tassos Kotsakis
andFedericaMarcolini gaveus unpublisheddata and sharedwith
us their opinion about the systematic position of some insular
rodents they are studying. Felisa Smith kindly provided us with a
new version of her mammalian body size database.
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No optimal body size in mammals
Global Ecology and Biogeography, 2010 Blackwell Publishing Ltd 9
BIOSKETCHES
Pasquale Raia is a post-doctoral research fellow at theDepartment of Earth Science, University of Naples
Federico II, and a member of the Center for
Evolutionary Ecology based at Rome III University. He
is interested in large mammal evolution, both at the
organismal and community levels, in response to
climate change and to the effect of ecological
interactions.
Francesco Carotenuto is a post-doctoral researchfellow at the Department of Earth Science University of
Naples. His research interests focus on Quaternary
mammal macroecology and biogeography.
Shai Meiri is a senior lecturer at the Department ofZoology, Tel Aviv University. He is interested in trait
evolution, the tempo and mode of evolution, the
evolutionary implications of biogeography, vertebrate
evolution and large wooden badgers.
Editor: Jos Alexandre F. Diniz-Filho
SUPPORTING INFORMATION
Additional Supporting Information may be found in the online
version of this article:
Appendix S1 Phylogenetic tree used for this study, along withextinct species included, their body size and source papers used
to ascertain both their size and phylogenetic position.
Appendix S2 Clade pairs partitioned in mainland and insulardaughter clades.
Appendix S3 Species belonging to clade pairs.
As a service to our authors and readers, this journal provides
supporting information supplied by the authors. Such materials
are peer-reviewed and may be reorganized for online delivery,
but are not copy-edited or typeset. Technical support issues
arising from supporting information (other than missing files)
should be addressed to the authors.
P. Raia et al.
Global Ecology and Biogeography, 2010 Blackwell Publishing Ltd10
1
APPENDIX S1
A. Extinct species included in this study B. References for size estimates and phylogenetic affinities for extinct species C. Phylogeny of all 4004 species used in the study
A. Extinct species included in the phylogenetic tree used for this study. For each species we report whether or not it was an insular endemic, its log 10 body mass (in grams), and the sources used to get body mass and phylogenetic data. Where the same source provided both size and phylogenetic data, the last column was left empty. Species insularity log
mass body mass reference Phylogeny reference
Capromeryx minor no 4.32 Brook & Bowman 2004 Janis & Manning 1998
Stockoceros conklingi no 4.72 Brook & Bowman 2004 Janis & Manning 1998
Tetrameryx shuleri no 4.79 Brook & Bowman 2004 Janis & Manning 1998
Capromeryx mexicana no 4.18 Smith et al. 2003 Janis & Manning 1998
Stockoceros onusrosagris no 4.74 Smith et al. 2003 Janis & Manning 1998
Bison antiquus no 6.01 Brook & Bowman 2004 Geraards 1992 Bison latifrons no 6.02 Brook & Bowman 2004 Geraards 1992 Bos primigenius no 5.97 Meloro et al., 2007 Geraards 1992 Bison priscus no 5.95 Smith et al. 2003 Geraards 1992 Pelorovis antiquus no 6 Smith et al. 2003 Geraards 1992 Bootherium bombifrons no 5.88 Brook & Bowman 2004 McDonald & Ray
1989 Euceratherium collinum no 5.7 Brook & Bowman 2004 McDonald & Ray
1989 Symbos cavifrons no 5.6 Smith et al. 2003 McDonald & Ray
1989 Megalotragus priscus no 5.3 Smith et al. 2003 Vrba 1979 Antidorcas australis no 4.6 Smith et al. 2003 placed with A.
marsupialis Antidorcas bondi no 4.53 Smith et al. 2003 placed with A.
marsupialis
2
Species insularity log mass
body mass reference Phylogeny reference
Oreamnos harringtoni no 4.88 Smith et al. 2003 placed with O. americanus
Camelus thomasi no 5.7 Smith et al. 2003 Feranec 2003 Hemiauchenia macrocephala no 5.04 Smith et al. 2003 Feranec 2003 Hemiauchenia paradoxa no 6 Smith et al. 2003 Feranec 2003 Palaeolama mirifica no 4.9 Smith et al. 2003 Feranec 2003 Dusicyon australis yes 4.699 Brook & Bowman 2004 Zrzavy & Rikankova
2003 Dusicyon avus no 4.11 Smith et al. 2003 Zrzavy & Rikankova
2003 Isolobodon portoricensis yes 3.1 Nowak 1999, It is a
conservative estimate as this species was probably larger than P. aedium
Woods et al. 2001
Plagiodontia ipnaeum yes 3.1 Nowak 1999, It is a conservative estimate as this species was probably larger than P. aedium
Woods et al. 2001
Hexolobodon phenax yes 3.75 Nowak 1999, It is suggested it was the size of Capromys pilorides
Woods et al. 2001
Isolobodon montanus yes 3.1 Nowak 1999, reportedly Isolobodon was the same size of Plagiodontia
Woods et al. 2001
Castoroides ohioensis no 5.18 Smith et al. 2003 placed sister to C. fiber
Praemegaceros cretensis yes 4.9 based on size ratio as reported in Raia & Meiri 2006
Croitor 2004, Raia & Meiri 2006
Praemegaceros ropalophorus yes 4.476 based on size ratio as reported in Raia & Meiri 2006
Croitor 2004, Raia & Meiri 2006
Praemegaceros cazioti yes 4.845 estimate from Burness et al. 2001
Croitor 2004, Raia & Meiri 2006
Megaloceros giganteus no 5.59 Meloro et al., 2007 Lister et al. 2005 Cervus astylodon yes 4.267 Matsumoto & Otsuka 2000; Based on the cubic
ratio of radius length as compared to Cervus nippon multiplied by the size of the latter. The morhotype used for comparison is G4 , which is the stratigraphically youngest.
3
Species insularity log mass
body mass reference Phylogeny reference
Cervalces scotti no 5.8 Smith et al. 2003 placed sister to Alces
Sinomegaceros yabei yes 5.589 van der Made & Tong 2008 state S. yabei is large, possibly as large as M. giganteus
Dasypus bellus no 4.65 Smith et al. 2003 Gaudin 2003, Vizcaino 2009
Holmesina septentrionalis no 5.4 Smith et al. 2003 Gaudin 2003, Vizcaino 2009
Holmesina occidentalis no 5.3 Smith et al. 2003 Gaudin 2003, Vizcaino 2009
Holmesina paulacoutoi no 5.1 Smith et al. 2003 Gaudin 2003, Vizcaino 2009
Pampatherium humboldtii no 5.18 Smith et al. 2003 Gaudin 2003, Vizcaino 2009
Pampatherium typum no 5.3 Smith et al. 2003 Gaudin 2003, Vizcaino 2009
Kraglievichia paranense no 4.653 Brook & Bowman 2004 Gaudin 2003, Vizcaino 2009
Sarcophilus laniarius no 4.08 Smith et al. 2003 placed sister to S. harrisi
Daubentonia robusta yes 4.7 Brook & Bowman 2004 name given to fossil remains of D. madagascarensis according to Wilson & Reeder (2003)
Diprotodon minor no 5.95 Smith et al. 2003 Black 2008 Diprotodon optatum no 6.18 Smith et al. 2003 Black 2008 Euryzygoma dunese no 5.7 Smith et al. 2003 Black 2008 Hulitherium thomasettii yes 5.14 Smith et al. 2003 Black 2008 Kolopsis watutense yes 5.48 Smith et al. 2003 Black 2008 Maokopia ronaldi yes 5 Smith et al. 2003 Black 2008 Nototherium mitchelli no 5.7 Smith et al. 2003 Black 2008 Zygomaturus trilobus no 5.88 Smith et al. 2003 Black 2008 Boromys offella yes 2.61 Turvey et al. 2007 Woods et al. 2001 Boromys torrei yes 2.29 Turvey et al. 2007 Woods et al. 2001 Brotomys voratus yes 2.86 Turvey et al. 2007 Woods et al. 2001 Heteropsomys insulans yes 3.34 Turvey et al. 2007 Woods et al. 2001 Elephas namadicus no 6.81 After measurements in
Maglio (1973) it is apparent E.antiquus & E.namadicus were the same body size
Thomas et al. 2000, Maglio 1973, Shoshani & Tassy 2005
4
Species insularity log mass
body mass reference Phylogeny reference
Elephas creutzburgi yes 6.57 based on size ratio as reported in Raia & Meiri 2006
Thomas et al. 2000, Maglio 1973, Shoshani & Tassy 2005
Elephas cypriotes yes 5.482 based on size ratio as reported in Raia & Meiri 2006
Thomas et al. 2000, Maglio 1973, Shoshani & Tassy 2005
Elephas mnaidriensis yes 6.272 based on size ratio as reported in Raia & Meiri 2006
Thomas et al. 2000, Maglio 1973, Shoshani & Tassy 2005
Mammuthus lamarmorae yes 5.812 based on size ratio as reported in Raia & Meiri 2006
Thomas et al. 2000, Maglio 1973, Shoshani & Tassy 2005
Palaeoloxodon naumanni yes 6.348 estimated by using regression equations in Roth (1990) & average shoulder height of 225 cm reported in Kondo et al. 2001
Thomas et al. 2000, Maglio 1973, Shoshani & Tassy 2005
Elephas antiquus no 6.81 Meloro et al., 2007 Thomas et al. 2000, Maglio 1973, Shoshani & Tassy 2005
Mammuthus columbi no 6.9 Smith et al. 2003 Thomas et al. 2000, Maglio 1973, Shoshani & Tassy 2005
Mammuthus imperator no 7 Smith et al. 2003 Thomas et al. 2000, Maglio 1973, Shoshani & Tassy 2005
Mammuthus primigenius no 6.74 Smith et al. 2003 Thomas et al. 2000, Maglio 1973, Shoshani & Tassy 2005
Mammuthus exilis yes 5.23 the cubed ratio of M.e. to Elephas falconeri's long bones multiplied by the estimated size of E.f.
Thomas et al. 2000, Maglio 1973, Shoshani & Tassy 2005
Onohippidium sp no 5.49 Stromberg 2006 Stromberg 2006 Equus hydruntinus no 5.32 Meloro et al., 2007 Burke et al. 2003 Hippidion principale no 5.71 Smith et al. 2003 Stromberg 2006 Hippidion saldiasi no 5.42 Smith et al. 2003 Stromberg 2006 Homotherium serum no 5.43 Brook & Bowman 2004 Slater & Van
Valkenburgh 2004
5
Species insularity log mass
body mass reference Phylogeny reference
Panthera atrox no 5.63 Brook & Bowman 2004 Slater & Van Valkenburgh 2005
Miracinonyx trumani no 4.94 Smith et al. 2003 Slater & Van Valkenburgh 2006
Smilodon fatalis no 5.64 Smith et al. 2003 Slater & Van Valkenburgh 2007
Smilodon populator no 5.65 Smith et al. 2003 Slater & Van Valkenburgh 2007
Giraffa gracilis no 5.93 Smith et al. 2003 placed sister to G. camaleopardalis
Chlamydotherium spp. no 5.24 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Glyptodon clavipes no 6.3 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Glyptodon reticulatus no 5.94 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Glyptotherium floridanum no 6.04 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Glyptotherium mexicanum no 6.04 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Lomaphorus spp. no 5.4 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Neosclerocalyptus spp. no 5.3 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Neothoracophorus depressus no 6.04 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Neothoracophorus elevatus no 5.9 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Panochthus tuberculatus no 6.03 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Plaxhaplous canaliculatus no 6.11 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
6
Species insularity log mass
body mass reference Phylogeny reference
Sclerocalyptus ornatus no 5.45 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Doedicurus clavicaudatus no 6.17 Smith et al. 2003 Gaudin 2003, Gaudin & Wible 2006, Vizcaino 2009
Cuvieronius hyodon no 6.623 Brook & Bowman 2004 Shoshani & Tassy 2005
Cuvieronius spp. no 6.7 Smith et al. 2003 Shoshani & Tassy 2005
Haplomastodon chimborazi no 6.78 Smith et al. 2003 Shoshani & Tassy 2005
Notiomastodon spp. no 6.79 Smith et al. 2003 Shoshani & Tassy 2005
Stegomastodon superbus no 6.88 Smith et al. 2003 Shoshani & Tassy 2005
Amblyrhiza inundata yes 4.97 McFarlane et al. 1998 Flemming & MacPhee 1996
Quemisia gravis yes 4.14 Nowak 1999, reportedly the same size of Elasmodontomys
Flemming & MacPhee 1996
Clidomys osborni yes 4.66 Nowak 1999. Weight estimate obtained as the cubic ratio of this species to C. pyloroides body size, the latter taken 5650 grams & 450 mm in head-body length
Flemming & MacPhee 1996
Clidomys parvus yes 4.36 Nowak 1999. Weight estimate obtained as the cubic ratio of this species to C. pyloroides body size, the latter taken 5650 grams & 450 mm in head-body length
Flemming & MacPhee 1996
Elasmodontomys obliquus yes 4.14 Smith et al. 2003 Flemming & MacPhee 1996
Hippopotamus madagascariensis yes 5.7 Brook & Bowman 2004 Boisserie 2005 Hippopotamus lemerlei yes 5.699 MOM-Mammals
Version 3.61 (current as of January 2007)
Boisserie 2005
Hippopotamus creutzburgi yes 5.741 Raia & Meiri 2006 Boisserie 2005 Hippopotamus pentlandi yes 6.182 Raia & Meiri 2006 Boisserie 2005
7
Species insularity log mass
body mass reference Phylogeny reference
Hippopotamus laloumena yes 5.98 Smith et al. 2003 Boisserie 2005 Neochoerus aesopi no 4.79 Prevosti & Vizcano
2006 Prado et al. 1998
Neochoerus pinckneyi no 4.85 Smith et al. 2003 Prado et al. 1998 Neochoerus sulcidens no 5.18 Smith et al. 2003 Prado et al. 1998 Archeolemur edwardsi yes 4.34 Brook & Bowman 2004 Orlando et al. 2008 Archeolemur majori yes 4.23 Brook & Bowman 2004 Orlando et al. 2008 Hadropithecus stenognathus yes 4.45 Brook & Bowman 2004 Orlando et al. 2008 Mesopropithecus dolichobrachion yes 4.08 Brook & Bowman 2004 Orlando et al. 2008 Mesopropithecus globiceps yes 4 Brook & Bowman 2004 Orlando et al. 2008 Mesopropithecus pithecoides yes 4.04 Brook & Bowman 2004 Orlando et al. 2008 Archaeoindris fontoynontii yes 5.3 Smith et al. 2003 Orlando et al. 2008 Pachylemur insignis yes 4 Brook & Bowman 2004 Orlando et al. 2008 Pachylemur jullyi yes 4.08 Brook & Bowman 2004 Orlando et al. 2008 Bohra paulae no 4.54 Brook & Bowman 2004 Flannery & Szalay
1982 (basal to other macropodinae)
Congruus gongruus no 4.6 Brook & Bowman 2004 probably basal to other macropodinae according to Prideaux 2004
Procoptodon goliath no 5.4 Brook & Bowman 2004 Prideaux 2004 Protemnodon nombe no 4.6 Brook & Bowman 2004 Prideaux 2004 Protemnodon tumbuna no 4.7 Brook & Bowman 2004 Prideaux 2004 Simosthenurus baileyi no 4.74 Brook & Bowman 2004 Prideaux 2004 Simosthenurus brachyselensis no 4.85 Brook & Bowman 2004 Prideaux 2004 Simosthenurus euryskaphus no 4.74 Brook & Bowman 2004 Prideaux 2004 Simosthenurus oreas no 5 Brook & Bowman 2004 Prideaux 2004 Sthenurus gilli no 4.48 Brook & Bowman 2004 Prideaux 2004 Procoptodon pusio no 4.88 Smith et al. 2003 Prideaux 2004 Procoptodon rapha no 5.18 Smith et al. 2003 Prideaux 2004 Protemnodon anak no 5 Smith et al. 2003 Prideaux 2004 Protemnodon brehus no 5 Smith et al. 2003 Prideaux 2004 Protemnodon hopei yes 4.87 Smith et al. 2003 Prideaux 2004 Protemnodon roechus no 4.95 Smith et al. 2003 Prideaux 2004 Simosthenurus brownei no 4.7 Smith et al. 2003 Prideaux 2004 Simosthenurus maddocki no 4.7 Smith et al. 2003 Prideaux 2004 Simosthenurus occidentalis no 4.7 Smith et al. 2003 Prideaux 2004 Sthenurus andersoni no 4.7 Smith et al. 2003 Prideaux 2004 Sthenurus atlas no 5.18 Smith et al. 2003 Prideaux 2004 Sthenurus stirlingi no 5.18 Smith et al. 2003 Prideaux 2004 Mammut americanum no 6.66 Smith et al. 2003 Yang et al. 1996 Megaladapis grandidieri yes 4.72 Smith et al. 2003 Orlando et al. 2008 Megaladapis madagascariensis yes 4.72 Smith et al. 2003 Orlando et al. 2008
8
Species insularity log mass
body mass reference Phylogeny reference
Megalonyx jeffersonii no 5.78 Smith et al. 2003 Gaudin 2004 Acratocnus odontrigonus yes 4.519 Brook & Bowman 2004 White & MacPhee
2001 Meizonyx salvadorensis no 5.778 Brook & Bowman 2004 White & MacPhee
2001 Neocnus comes yes 3.778 Brook & Bowman 2004 White & MacPhee
2001 Parocnus serus yes 4.845 Brook & Bowman 2004 White & MacPhee
2001 Megalocnus rodens yes 5.176 estimate from Burness
et al. 2001 White & MacPhee 2001
Megalocnus zile yes 5.176 estimate from Burness et al. 2001
White & MacPhee 2001
Acratocnus antillensis yes 4.631 Estimated from post-cranial material in Arredondo & Arredondo 2000
White & MacPhee 2001
Neocnus gliriformis yes 4.078 Estimated from post-cranial material in Arredondo & Arredondo 2000
White & MacPhee 2001
Neocnus major yes 4.164 Estimated from post-cranial material in Arredondo & Arredondo 2000
White & MacPhee 2001
Parocnus browni yes 4.942 Estimated from post-cranial material in Arredondo & Arredondo 2000
White & MacPhee 2001
Acratocnus simorhincus yes 4.176 Rega et al. 2002 White & MacPhee 2001
Valgipes spp. no 5.3 Smith et al. 2003 White & MacPhee 2001
Nothrotheriops shastensis no 5.79 Brook & Bowman 2004 Gaudin 2004 Eremotherium laurillardi no 5.9 Smith et al. 2003 Gaudin 2004 Eremotherium rusconii no 6.54 Smith et al. 2003 Gaudin 2004 Megatherium americanum no 6.8 Smith et al. 2003 Gaudin 2004 Nothropus spp. no 5 Smith et al. 2003 Gaudin 2004 Nothrotherium spp. no 5.18 Smith et al. 2003 Gaudin 2004 Ocnopus spp. no 5.48 Smith et al. 2003 Gaudin 2004 Paramegatherium spp. no 6.54 Smith et al. 2003 Gaudin 2004
9
Species insularity log mass
body mass reference Phylogeny reference
Terricola melitensis yes 1.339 A. Kotsakis, pers. Comm. Estimated by multiplying upper tooth row ratio per body weight of M. duodecimcustatus (data kindly provided by F. Marcolini)
Meriones malatestai yes 2.1 A. Kotsakis, pers. Comm. Estimated by multiplying upper tooth row ratio per body weight of M. tristrami
Oryzomys nelsoni yes 2.017 based on comparison with O. palustris. Both species measurements were taken from MAMMALIAN SPECIES accounts
Mus lopadusae yes 1.611 Burgio & Catalisano 1994 Nesoryzomys swarthi yes 1.97 Dowler et al. 2000 Mus minotaurus yes 1.526 Federica Marcolini, pers. Comm. Rhagamys orthodon yes 1.801 Gliozzi et al. 1984. based on m1 length of
Apodemus flavicollis
Spelaeomys florensis yes 2.98 Musser 1981, . Estimated multiplying the cubic ratio of m1-3 length with that of Rattus fuscipes
Paulamys naso yes 2.09 Nowak 1999 Neotoma bunkeri yes 2.57 Smith et al. 2003 Likely conspecific
with N. lepida according to Wilson & Reeder (2003)
Uromys imperator yes 3 Smith et al. 2003 placed in the U. rex species group according to Wilson & Reeder (2003)
Papagomys theodorverhoeveni yes 3 Zijlstra et al. 2008, Estimated for the cubic ratio of m1 length as compared with P. armandvillei multiplied by the size of the latter
Lutrogale cretensis yes 4.041 estimate from Burness et al. 2001
placed in the European otter species group
Sardolutra ichnusae yes 4.031 Malatesta 1977 Algarolutra majori yes 4.156 Malatesta et al. 1986
Lutra euxena yes 3.944 Willemsen 1992. Cubic ratio of m1 lengths as compared with L. lutra, multiplied by the weight of the latter
10
Species insularity log mass
body mass reference Phylogeny reference
Lutra trinacrie yes 4.051 Willemsen 1992. Cubic ratio of m1 lengths as compared with L. lutra, multiplied by the weight of the latter
Megalenhydris barbaricina yes 4.443 Willemsen 1992. Cubic ratio of m1 lengths as compared with L. lutra, multiplied by the weight of the latter
Glossotherium harlani no 6.3 Brook & Bowman 2004 Gaudin 2004 Glossotherium myloides no 6.08 Smith et al. 2003 Gaudin 2004 Glossotherium robustum no 6.23 Smith et al. 2003 Gaudin 2004 Lestodon armatus no 6.53 Smith et al. 2003 Gaudin 2004 Mylodon listai no 6 Smith et al. 2003 Gaudin 2004 Scelidodon spp. no 6 Smith et al. 2003 Gaudin 2004 Scelidotherium leptocephalum no 6.05 Smith et al. 2003 Gaudin 2004 Eliomys morpheus yes 2.4 Alcover et al. 2000 Muscardinus malatestai yes 1.633 The cubed ratio of M1 length compared with
M. avellanarius multplied by the size of the latter. Measurements in Gliozzi 1995
Nesophontes edithae yes 2.32 Turvey et al. 2007 Asher 1999 Nesophontes hypomicrus yes 1.36 Turvey et al. 2007 Asher 1999 Nesophontes micrus yes 1.66 Turvey et al. 2007 Asher 1999 Nesophontes paramicrus yes 1.67 Turvey et al. 2007 Asher 1999 Nesophontes zamicrus yes 1 Turvey et al. 2007 Asher 1999 Babakotia radofilai yes 4.18 Smith et al. 2003 Orlando et al. 2008 Palaeopropithecus ingens yes 4.68 Smith et al. 2003 Orlando et al. 2008 Palaeopropithecus maximus yes 4.68 Smith et al. 2003 Orlando et al. 2008 Palorchestes azael no 5.7 Brook & Bowman 2004 Black 2008 Palorchestes parvus no 5 Smith et al. 2003 Black 2008 Phascolarctos stirtoni no 4.15 Smith et al. 2003 placed sister to P.
cinereus Coelodonta antiquitatis no 6.46 Brook & Bowman 2004 Cerdeno 1995 Solenodon arredondoi yes 2.97 Turvey et al. 2007 Nesiotites similis yes 1.38 A. Kotsakis, pers. Comm. Estimated by
comparing upper tooth row length with that of A. hidalgo
Asoriculus hidalgo yes 1.3 Alcover et al. 2000 Stegodon orientalis no 6.3 estimate based on Van den Bergh ( 1997) size
estimate of Javan S. trigonocephalus at 1.7 tons. S.t. is said to be slightly smaller than S. orientalis
11
Species insularity log mass
body mass reference Phylogeny reference
Stegodon sp. yes 5.54 Van den Bergh, G.D. 1997. The Late Neogene. No measurements are available for dwarf Stegodon associated to Homo floresiensis. Possibly late-Pleistocene, truly dwarf Stegodon are known from Sumba & Timor. We used the mass estimate calculated on S. sampoensis
Metridiochoerus andrewsi no 5.18 Brook & Bowman 2004 White & Harris 1977 Metridiochoerus compactus no 5.15 Smith et al. 2003 White & Harris 1977 Zaglossus hacketti no 4.48 Smith et al. 2003 sister to Z. bruijni Platygonus compressus no 5.04 Smith et al. 2003 Wetzel et al. 1975 Arctodus pristinus no 5.48 Smith et al. 2003 McLellan & Rainer
2004 Arctodus simus no 5.86 Smith et al. 2003 McLellan & Rainer
2004 Tremarctos floridanus no 5.18 Smith et al. 2003 McLellan & Rainer
2004 Microtus henseli yes 1.301 Gliozzi et al. 1984, based on M1 of M. savii
Phanourios minor yes 5.58 Raia & Meiri 2006 placed sister to H. amphibious
Megatapirus augustus no 5.76 Tong 2005
12
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C. The phylogeny used in the study (((Tachyglossus_aculeatus:10.6,(Zaglossus_bruijni:5.3,Zaglossus_hacketti:5.3)4008:5.3)4007:53,Ornithorhynchus_anatinus:63.6)4006:102.800004,(((((((((((((Anomalurus_beecrofti:23.6,Anomalurus_derbianus:23.6,Anomalurus_pelii:23.6,Anomalurus_pusillus:23.6)4021:7.6,(Zenkerella_insignis:18.7,(Idiurus_macrotis:12.9,Idiurus_zenkeri:12.9)4023:5.8)4022:12.5)4020:23.7,Pedetes_capensis:54.9)4019:22.5,(((Allactaga_elater:24.5,Allactaga_bullata:24.5,Allactaga_euphratica:24.5,Allactaga_hotsoni:24.5,Allactaga_major:24.5,Allactaga_severtzovi:24.5,Allactaga_sibirica:24.5,Allactaga_tetradactyla:24.5)4026:14.5,((Paradipus_ctenodactylus:38.5,Euchoreutes_naso:38.5,(Sicista_betulina:26.7,Sicista_tianshanica:26.7,Sicista_caucasica:26.7,Sicista_caudata:26.7,Sicista_concolor:26.7,Sicista_napaea:26.7,Sicista_strandi:26.7,Sicista_subtilis:26.7)4029:11.8,(Dipus_sagitta:11.8,Eremodipus_lichtensteini:11.8,(Jaculus_jaculus:11.5,Jaculus_blanfordi:11.5,Jaculus_orientalis:11.5)4031:0.3,(Stylodipus_andrewsi:9.2,Stylodipus_telum:9.2)4032:2.6)4030:26.7,(Napaeozapus_insignis:11.9,Eozapus_setchuanus:11.9,(Zapus_hudsonius:10.1,Zapus_princeps:10.1,Zapus_trinotatus:10.1)4034:1.8)4033:26.6,(Cardiocranius_paradoxus:20.2,(Salpingotus_crassicauda:18.7,Salpingotus_kozlovi:18.7,Salpingotus_michaelis:18.7)4036:1.5)4035:18.3)4028:0.3,Pygeretmus_pumilio:38.8)4027:0.2)4025:31.3,(((Abditomys_latidens:14.3,(Bullimus_bagobus:4.7,Bullimus_luzonicus:4.7)4040:9.6)4039:14.8,(((Acomys_cahirinus:5.3,Acomys_ignitus:5.3,Acomys_kempi:5.3,Acomys_minous:5.3,Acomys_percivali:5.3,Acomys_russatus:5.3,Acomys_spinosissimus:5.3,Acomys_subspinosus:5.3,Acomys_wilsoni:5.3)4043:7,Acomys_louisae:12.3)4042:0.2,(Lophuromys_flavopunctatus:5.8,Lophuromys_melanonyx:5.8,Lophuromys_nudicaudus:5.8,Lophuromys_rahmi:5.8,Lophuromys_sikapusi:5.8,Lophuromys_woosnami:5.8)4044:6.7,Uranomys_ruddi:12.5)4041:16.6,(Aethomys_chrysophilus:19.9,Aethomys_granti:19.9,Aethomys_hindei:19.9,Aethomys_kaiseri:19.9,Aethomys_namaquensis:19.9,Aethomys_nyikae:19.9)4045:9.2,(Anisomys_imitator:26.5,(Chiruromys_forbesi:7.4,Chiruromys_lamia:7.4,Chiruromys_vates:7.4)4047:19.1,Coccymys_ruemmleri:26.5,Crossomys_moncktoni:26.5,(Hyomys_dammermani:4.7,Hyomys_goliath:4.7)4048:21.8,((Leptomys_elegans:6.6,Leptomys_ernstmayri:6.6)4050:9,Lorentzimys_nouhuysi:15.6,(Mayermys_germani:4.2,Mayermys_ellermani:4.2,Neohydromys_fuscus:4.2)4051:11.4,(Paraleptomys_rufilatus:4.2,Paraleptomys_wilhelmina:4.2)4052:11.4,(Pseudohydromys_murinus:4.2,Pseudohydromys_occidentalis:4.2)4053:11.4)4049:10.9,Macruromys_major:26.5,(Mallomys_aroaensis:9.4,Mallomys_gunung:9.4,Mallomys_istapantap:9.4,Mallomys_rothschildi:9.4)4054:17.1,Microhydromys_richardsoni:26.5,Parahydromys_asper:26.5,(Pogonomelomys_mayeri:7.4,Pogonomelomys_sevia:7.4)4055:19.1,(Solomys_ponceleti:10.9,Solomys_salebrosus:10.9)4056:15.6,Xenuromys_barbatus:26.5)4046:2.6,((((Apodemus_argenteus:11.1,Apodemus_draco:11.1,Apodemus_gurkha:11.1,Apodemus_latronum:11.1,Apodemus_peninsulae:11.1,Apodemus_semotus:11.1,Apodemus_speciosus:11.1)4060:5.5,(Apodemus_agrarius:3,Apodemus_chevrieri:3)4061:13.6,Apodemus_mystacinus:16.6,(Apodemus_alpicola:4.9,Apodemus_flavicollis:4.9,Apodemus_fulvipectus:4.9,Apodemus_hermonensis:4.9,Apodemus_sylvaticus:4.9,Apodemus_uralensis:4.9)4062:11.7)4059:3.4,Rhagamys_orthodon:20)4058:3.1,(Tokudaia_muenninki:4.8,Tokudaia_osime
17
nsis:4.8)4063:18.3)4057:6,((Apomys_abrae:3.9,Apomys_datae:3.9,Apomys_hylocoetes:3.9,Apomys_insignis:3.9,Apomys_littoralis:3.9,Apomys_microdon:3.9,Apomys_musculus:3.9,Apomys_sacobianus:3.9)4065:5,(((Archboldomys_luzonensis:3.6,(Crunomys_celebensis:2.9,Crunomys_fallax:2.9,Crunomys_melanius:2.9)4069:0.7)4068:0.1,(Celaenomys_silaceus:2.9,(Chrotomys_gonzalesi:0.8,Chrotomys_mindorensis:0.8,Chrotomys_whiteheadi:0.8)4071:2.1)4070:0.8)4067:0.2,(Rhynchomys_isarogensis:2.6,Rhynchomys_soricoides:2.6)4072:1.3)4066:5)4064:20.2,(Arvicanthis_abyssinicus:10.9,Arvicanthis_blicki:10.9,Arvicanthis_nairobae:10.9,Arvicanthis_niloticus:10.9)4073:18.2,(Bandicota_bengalensis:8.1,Bandicota_indica:8.1,Bandicota_savilei:8.1)4074:21,((Batomys_dentatus:7.4,Batomys_granti:7.4,Batomys_salomonseni:7.4)4076:7.9,(Crateromys_australis:7.4,Crateromys_paulus:7.4,Crateromys_schadenbergi:7.4)4077:7.9)4075:13.8,(Berylmys_berdmorei:10.2,Berylmys_bowersi:10.2,Berylmys_mackenziei:10.2,Berylmys_manipulus:10.2)4078:18.9,((Bunomys_andrewsi:14.3,Bunomys_chrysocomus:14.3,Bunomys_coelestis:14.3,Bunomys_fratrorum:14.3,Bunomys_heinrichi:14.3,Bunomys_penitus:14.3,Bunomys_prolatus:14.3)4080:7.4,Paulamys_naso:21.7)4