Raia Et Al. 2010 One Size Does Not Fit All

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.


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



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.


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.



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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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.


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


Praemegaceros cazioti yes 4.845 estimate from Burness et al. 2001


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



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



Elephas creutzburgi yes 6.57 based on size ratio as reported in Raia & Meiri 2006


Elephas cypriotes yes 5.482 based on size ratio as reported in Raia & Meiri 2006


Elephas mnaidriensis yes 6.272 based on size ratio as reported in Raia & Meiri 2006


Mammuthus lamarmorae yes 5.812 based on size ratio as reported in Raia & Meiri 2006


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


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.


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



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



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


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


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



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



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


Neocnus gliriformis yes 4.078 Estimated from post-cranial material in Arredondo & Arredondo 2000


Neocnus major yes 4.164 Estimated from post-cranial material in Arredondo & Arredondo 2000


Parocnus browni yes 4.942 Estimated from post-cranial material in Arredondo & Arredondo 2000


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



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



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



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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62 Zrzavy, J. and Rikankova, V. 2004. Phylogeny of Recent Canidae (Mammalia, Carnivora): relative reliability and utility of morphological and molecular datasets. Zoologica Scripta, 33, 311333.

16

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

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