Integration of global fossil and modern biodiversity data ...

ORIGINALARTICLE

Integration of global fossil and modernbiodiversity data reveals dynamism andstasis in ant macroecological patternsBenoit Gu�enard1,2*, Vincent Perrichot3 and Evan P. Economo1,4

1Okinawa Institute of Science and Technology

Graduate University, Onna-son, Okinawa

904-0495, Japan, 2School of Biological

Sciences, The University of Hong Kong, Pok

Fu Lam Road, Hong Kong SAR, China,3CNRS UMR 6118 G�eosciences, Universit�e de

Rennes 1, Rennes Cedex 35042, France,4Department of Ecology and Evolutionary

Biology, University of Michigan, Ann Arbor

MI 48104, USA

*Correspondence: Benoit Gu�enard, School of

Biological Sciences, The University of Hong

Kong, Pok Fu Lam Road, Hong Kong SAR,

China.

E-mail: [email protected]

ABSTRACT

Aim We investigate the dynamics of ant biodiversity patterns and community

structure from the Eocene until the present. Our goal is to empirically test

hypotheses regarding the similarities in composition and structure of fossil and

modern ant faunas to understand the historical processes influencing modern

global biodiversity patterns.

Location Global.

Methods We integrated two recently developed databases of the geographical

distributions of fossil and modern ant genera and analysed the evolution of

diversity and composition since the Eocene, c. 55 Ma. We assembled a third new

database on community structure of ants, and used it to compare community

structure of fossil assemblages with modern communities in different bioregions.

Results The analyses of generic composition and community structure were

congruent, supporting a strong affinity between the Western Palaearctic ant

fauna and modern Indomalayan and Australasian assemblages, and of a wide-

spread Holarctic ant palaeofauna, and affinity between fossil Caribbean and

modern Neotropical faunas. In addition, neither generic composition nor com-

munity structure of fossil assemblages showed evidence of taphonomic bias

towards arboreal taxa.

Main Conclusions The aggregated fossil record reveals the dynamic nature of

macroecological patterns in ant biodiversity during the Cenozoic, with conti-

nental-scale generic extinctions common and important in shaping modern ant

assemblages. Our results suggest major compositional changes for the Western

Palaearctic bioregion and the Caribbean bioregions. The ant palaeofauna of the

Western Palaearctic bioregion was then very similar in composition and struc-

ture to the one observed now in the Indomalayan bioregion, supporting the

notion that modern-day ‘tropical’ biomes were historically much more wide-

spread, while temperate biotas are more recently evolved. Our results under-

score the importance of a palaeogeographical perspective in understanding

modern-day macroecological patterns.

Keywords

Cenozoic, community structure, extinctions, Formicidae, fossils, generic

diversity, global diversity patterns, historical biogeography, neofauna, palaeo-

fauna.

INTRODUCTION

Biologists have long strived to understand the macroecologi-

cal patterns such as the latitudinal diversity gradient and

the distribution of Earth’s biomes. In recent years, these

efforts have been bolstered by new informatic tools allowing

for aggregations of vast amounts of data. As noted by

others (Manchester et al., 2009; Jones & Safi, 2011), the

large-scale biodiversity patterns we observe today are an

outcome of a long and dynamic evolutionary history driven

ª 2015 John Wiley & Sons Ltd http://wileyonlinelibrary.com/journal/jbi 1doi:10.1111/jbi.12614

Journal of Biogeography (J. Biogeogr.) (2015)

by climatological, geological and ecological shifts. The

aggregation and synthesis of fossil data is critical for resolv-

ing the temporal dimension of biodiversity dynamics and

understand the processes driving patterns.

While data syntheses have advanced rapidly for vertebrates

and plants, invertebrates lag behind despite constituting the

large majority of the global diversity. Among invertebrates,

ants are one of the most ecologically dominant and well-

studied groups. Understanding the mechanisms and the

pathways by which this group achieved such ecological and

evolutionary success represents a major endeavour. Recently,

a global database on the distribution of modern ant genera

was compiled revealing large-scale patterns in richness and

composition (Gu�enard et al., 2012). Work on fossil ants has

mainly focused on taxonomic analysis of individual speci-

mens (e.g. Baroni Urbani, 1980; Perrichot et al., 2008a;

Dlussky & Wedmann, 2012). Only a few synthetic analyses

exist on particular deposits with a rich ant palaeocommunity,

such as Baltic and Dominican ambers (Wheeler, 1915; Wil-

son, 1985; Dlussky & Rasnitsyn, 2009; Dlussky & Putyatina,

2014). Recently, a world catalogue of fossil ants was com-

piled (Perrichot, 2014), and the updated global fossil record

was reviewed (LaPolla et al., 2013). However, our macro-

scopic view of ant biodiversity, across all genera and across

the globe, remains firmly rooted in the present.

Here, for the first time, we assemble a comprehensive

global database of ant genera known both from fossil and

modern species, providing relevant historical context to the

patterns observed in extant generic distributions. We evaluate

the extent to which historical assemblages of genera resemble

modern faunas, both in the same and in distant geographical

regions. We use these data on generic composition to evalu-

ate biogeographical hypotheses developed to explain modern

ant composition patterns.

While generic composition varies in informative ways

across major bioregions, the distribution of diversity at

higher taxonomic levels also varies in important ways that

are not captured by the presence–absence of genera. For

example, compared with faunas in other continents, both

local and regional ant faunas of Australia are much more

dominated by species of the subfamily Dolichoderinae than

other regions. These broad differences in community organi-

zation also help us interpret the organization of historical

ant assemblages. Likewise fossil records may be biased

towards ants living in certain habitats (forest, open, etc.) and

in different ecological strata (arboreal, epigaeic, hypogaeic)

as noted for instance by Dlussky et al. (2009) on the domi-

nance of arboreal ant individuals in the Oligocene Bembridge

Marls deposits.

To compare community organization of fossil and modern

assemblages, we compiled a global database of modern ant

community structure from all major biomes and habitat

types, as characterized by the distribution of species richness

across subfamilies. Using this data set, we evaluated whether

the organization of fossil communities resembles any mod-

ern-day faunas, and if there is evidence of a taphonomic bias

towards certain habitats or strata. We first review large-scale

ant biogeography to establish some context and hypotheses

for our analysis.

Background and hypotheses

Recent work on ant evolution including molecular phyloge-

netic analyses (Brady et al., 2006; Moreau et al., 2006; Mor-

eau & Bell, 2013), diversification hypotheses (Wilson &

H€olldobler, 2005; Moreau et al., 2006; Perrichot et al.,

2008b), and new description of ancient fossils nearly 100 Ma

old (Nel et al., 2004; Perrichot et al., 2008a; Barden & Gri-

maldi, 2012), indicate that all the genus-level taxa known

from the first 50–60 Myr following their first appearance are

now extinct. The rise of the modern ant fauna is a more

recent phenomenon occurring probably during the early

Cenozoic and continuing until the present (Brady et al.,

2006; Moreau et al., 2006; LaPolla et al., 2013).

The Fossil West Palaearctic–Modern Indomalayan-

Australasian Affinity

One of the pioneering -and still most impressive- studies on

fossil ants to-date was conducted by Wheeler (1915) on 9527

ant specimens from Eocene (37–42 Ma) Baltic amber.

Wheeler noted then that modern American and African ele-

ments are nearly completely missing from Baltic amber, and

he described this ant palaeofauna as a ‘mixture of what at

the present day we are able to recognize as at least four dif-

ferent faunas, the Palaearctic, the Indian, the Malayan and

the Australian, with a little more than ⅓ of the genera and

nearly ½ of the species Palaearctic and the reminder belong-

ing to Indomalayan and Australian types’ (Wheeler, 1915, p.

12). Wheeler’s observations and conclusions followed those

of Emery (1892), who previously noticed the strong affinity

of much smaller sample of Sicilian amber ant fossils with the

modern Indo-Australian fauna. However, these authors ver-

bally described patterns but did not test them quantitatively.

Furthermore, as those original studies, subsequent work has

enriched our understanding of both the fossil and modern

ant faunal distributions, with studies on new deposits avail-

able (Europe: Dlussky & Rasnitsyn, 2009; Dlussky & Wed-

mann, 2012; Dominican amber: Wilson, 1985; Baroni

Urbani, 1995; Chiapas: Sol�orzano Kraemer, 2007), and

improvements in taxonomy of both modern and fossil taxa

(Ward, 2007). The aggregation of the most recent data for

both the fossil and modern ant distributions (Gu�enard et al.,

2010; Perrichot, 2014) allow now for more rigorous compar-

ison of the affinities between those different assemblages.

The East Palaearctic–East Nearctic Disjunction

For over 150 years, plant ecologists have noticed strong simi-

larities between plant communities found in East Asia and

East North America, both sharing similar genera, while those

were absent from Europe or West America (Wen, 1999).

Journal of Biogeographyª 2015 John Wiley & Sons Ltd

2

B. Gu�enard et al.

Those observations led to the hypothesis that during most of

the Cenozoic a large mixed mesophytic forest covered the

Northern Hemisphere (Tiffney, 1985; Manchester, 1999;

Greenwood et al., 2005), with the North Atlantic and Bering

land bridges connecting Eurasia and North America facilitat-

ing transcontinental dispersal of plant and animal lineages.

By the late Tertiary and during the Quaternary, major cool-

ing events provoked massive extinctions in Europe and part

of the Nearctic bioregion (Tiffney, 1985) and closed overland

dispersal routes. Despite support from numerous plants and

vertebrate studies (Tiffney, 1985; Manchester, 1999; San-

martin et al., 2001), this hypothesis has received less atten-

tion for entire communities of insects groups (but see Von

Dohlen et al., 2002). If valid, we predict that the fossil ant

assemblages known from the Holarctic bioregion (East

Palaearctic, West Palaearctic and Nearctic bioregions) should

be aligned with the current faunas of the Eastern Asian and

Eastern Nearctic bioregions. Alternatively, regions that were

affected by extinctions should present limited affinities

between their Palaeo- and Neofauna.

Similarity between the current Neotropical bioregion and

the Miocene Caribbean fauna

A comparison of the Hispaniolan modern fauna and palaeo-

fauna conducted by Wilson (1985) showed the similarity of

the past Dominican amber ant fauna with the current ant

fauna of the Neotropical bioregion. A similar pattern with

moderate similarity between modern and palaeo-Hispaniolan

fauna, and strong similarities between palaeo-Hispaniolan

and modern South and North American fauna has been

observed in several groups of insects (Sol�orzano Kraemer,

2007) and spiders (Penney, 2008). In contrast, Baroni Urbani

and de Andrade highlighted examples of Dominican fossils

that had strong affinities with extant Old World ant fauna,

in particular the current Australasian taxa (Baroni Urbani,

1980, 1995; Baroni Urbani & de Andrade, 2003), a pattern

also observed occasionally in other insect groups (Grimaldi

et al., 2013). We use our data to evaluate the extent to which

the Caribbean ant palaeofauna is related to the Neotropical

fauna versus faunas of other bioregions.

Community organization across bioregions

In addition to the presence–absence of genera, other aspects

of community organization exhibit biogeographical variation.

In ant assemblages, the relative proportion of the species

from the different subfamilies varies at both local and large

scales, and across habitat strata. In tropical forests, stratifica-

tion among microhabitats are well documented, with few

overlaps between species inhabiting the arboreal stratum and

those living on the ground surface or deeper in the leaf litter

(Wilson, 1959; Br€uhl et al., 1998). At a continental-scale,

Ward (2000) briefly described major geographical differences

in how species richness is partitioned among subfamilies in

leaf-litter assemblages.

However, such patterns may be influenced by taphonomic

bias. Several studies of European fossil deposits found that

the abundance and richness of ant species from the subfam-

ily Dolichoderinae in ant communities was much higher

than usually observed in modern communities (Wheeler,

1915; Dlussky & Rasnitsyn, 2002; LaPolla et al., 2013). This

led these authors to suggest that because modern Doli-

choderinae are most commonly arboreal, the taphonomy of

amber fossilization could be biased towards arboreal species.

However, more general palaeoecological studies have shown

that a significant fraction of organisms embedded in amber

were representatives of the litter and low vegetation of the

forest (Henwood, 1993; Perrichot, 2004). Thus, it is possible

that it is the preponderance of Dolichoderines on other sub-

family-level taxa which (according to the hypothesis

described above) may reflect similarities to modern Indoma-

layan and Australasian faunas. In contrast, if the taphonomy

of ant fossils is biased towards arboreal taxa, then we should

find greater affinities with community structure found in

contemporary arboreal communities, but not particularly

allied with a given bioregion.

To summarize, in this study we use three recently devel-

oped databases to address the following goals. (1) By com-

paring past and modern distributions of ant genera, we

explore the validity of multiple biogeographical hypotheses

developed to explain modern ant composition patterns,

namely (i) the fossil West Palaearctic ant fauna affinity with

the modern Indomalayan-Australasian fauna, (ii) the East

Nearctic–East Palaearctic disjunction, and (iii) the fossil Car-

ibbean affinity with the modern Neotropical fauna. (2) We

study the relative species richness of ant subfamilies within

communities present in multiple modern bioregions and

habitats in order to characterize the structural affinities of

European fossil assemblages of the Palaeogene period. (3)

We test the hypothesis that the generic representation and

community structure within the European fossil record is

biased towards an over-representation of arboreal species.

METHODS

Fossil genera database

We focused on ant genera known from both the fossil record

and the modern ant fauna, as these provide information

about affinities between historical and extant faunas. Fossils

of ants were assigned to specific deposits representing a given

area and a specific geological period. We assigned each

deposit to a bioregion in order to compare generic fossil and

modern ant records. Table S1 (see Appendix S2 in Support-

ing Information) presents the main geological deposits where

ants have been collected and the bioregions and periods to

which they were assigned. Controversial fossil records were

excluded (see details in Appendix S1).

Nine bioregions were considered in this study: the West

Palaearctic, East Palaearctic, Indomalayan, Australasian,

Afrotropical, Malagasy, Nearctic, Caribbean and the


3

Historical biogeography of ants

Neotropical bioregion (see Appendix S1). Figure S1 in

Appendix S2 presents how the political regions used

in Gu�enard et al. (2012) were assigned to each bioregion. In

order to evaluate the similarity between the ant fossil fauna

of the Dominican amber with other bioregions, we consid-

ered the Caribbean as its own bioregion. This choice was

motivated by the results of previous studies on the dissimi-

larity between the modern and the Dominican ant palaeo-

fauna (Wilson, 1985; Baroni Urbani, 1995).

Community database

To compare the community structure of fossil and modern

ant assemblages, we built a database from 1060 publica-

tions, providing species composition at local and regional

scales. Species richness for each of the 16 extant ant sub-

families was extracted, and the relative proportion of spe-

cies richness (number of species in a subfamily/total species

richness collected) was calculated. Other relevant data, such

as spatial coordinates, elevation, habitat type, collection

techniques and target strata (hypogaeic, epigaeic, arboreal

or any combination of those) were also noted. After filter-

ing the sites due to certain criteria (see Appendix S1), 1736

sites representative of modern communities were used for

the analysis. Details on the number of sites within each

bioregion and for each habitat are given in Table S2 in

Appendix S2.

Modern ant assemblages were compared to the fossil

assemblages of the Baltic, Bitterfeld, Rovno and Scandinavian

amber deposits compiled by Dlussky & Rasnitsyn (2009) rep-

resenting over 16,700 inclusions (respectively Baltic = 14,915,

Bitterfeld = 1039; Rovno = 501; and Scandinavian = 271).

Similar species-level data were not readily available for other

deposits. For this analysis, all taxa, independently of their

status of extinct or extant, were included in the fossil ant

community assessment.

Analyses

To evaluate affinities between historical and modern biore-

gions, we calculated the fraction of genera present in each

fossil deposit that are also present in the extant fauna of each

bioregion, considering only those genera from the fossil

record that are still extant somewhere in the world.

To evaluate taphonomic bias based on generic composi-

tion, we classified the extant genera known from the fossil

records into one of the following categories: arboreal, epi-

gaeic, hypogaeic, arboreal + epigaeic, epigaeic + hypogaeic,

arboreal + epigaeic + hypogaeic (no genera were classified as

arboreal + hypogaeic). Furthermore, we considered if the

fossil specimens for each genus were known as worker (non-

flying individual) or alate (flying gyne or male). This is an

important consideration as most compression fossils

(through sedimentation in water) are represented by alate

individuals and many hypogaeic species are over-represented

by alate individuals trapped in resin during mating flights

(Dlussky & Rasnitsyn, 2009). Our null hypothesis considers

that all else equal, the proportion of genera known from

each category (arboreal, epigaeic, hypogaeic) should be equal

if unbiased towards one category. The tested hypothesis here

is that the proportion of genera known to live within each

stratum does not show a bias towards a specialized habitat

life style. This represents the simplest hypothesis to show

that all strata-inhabitants could be preserved in fossilization

processes. We used a chi-square analysis to test this hypothe-

sis both including and excluding alate individuals from the

records.

Using our databases of fossil and extant community com-

position, we asked if fossil assemblages resemble any extant

communities somewhere on the globe (bioregion, habitat,

and ecological stratum) based on the distribution of richness

across subfamilies. We took two approaches to answering

this question. First, we used nonmetric multidimensional

scaling (NMDS) to characterize the variability in community

composition within and across bioregions, and place the fos-

sil communities in this context. Second, we used a multino-

mial regression approach to statistically assess which modern

faunas most closely resemble the fossil data.

Ordinations

Nonmetric multidimensional scaling and related ordination

techniques take high dimensional data and arrange it in a

low-dimensional form where distance between points in the

low dimensional space is representative of distance in the

higher dimensional space. We performed this analysis to

visualize where the fossil communities fall in the context of

the extant communities. We first calculated the proportion

of the species richness found in each subfamily. We then cal-

culated the Euclidean distance between all community pairs,

and performed a 2-dimensional NMDS using Matlab and

the sum of squared distance as the stress criterion.

Multinomial logistic regression analysis

We used multinomial logistic regression to fit a model for

community composition (distribution of species richness

across subfamilies), using bioregion, habitat type, and stra-

tum as predictor variables. Then, using the models fitted to

extant communities, we ask which one best predicts the

observed composition of the fossil assemblage. The model

predicts the vector of frequencies across each subfamily, and

a community of k species is treated as a multinomial draw k

times from that generating distribution. This multinomial

regression approach accounts for the noise introduced by

sampling effects in low richness sites (due either to actual

low richness or undersampling), thus we could include low

richness sites giving us a total of 1736 communities in the

analysis.

We coded the nine bioregions and four habitat types as

binary indicator variables in the analysis. For the three strata

(ground, arboreal, hypogaeic), some sampling efforts focused


4


on more than one stratum. For those we used a dummy

coefficient of either 1/2 if two of the three were used or 1/3

if all three were used. In this way we let some community

samples be a mixture of different strata. After fitting the

model to the extant community data set, we used the fitted

model to predict a probability vector for each combination

of predictors (each biome times each habitat type times all

combinations of strata). We then calculated the likelihood of

each multinomial model given each set of fossil data from

the four deposits and all fossil data combined.

RESULTS

We gathered 227 unique fossil per period records for 74 ant

genera spread over 10 subfamilies, which represent c. 23% of

all modern ant genera and 63% of modern ant subfamilies.

Based on the age estimation given to each fossil deposit (see

Table S1 in Appendix S2), 35, 25 and 64 genera are known

respectively from the Eocene, the Oligocene, and the Mio-

cene (see Fig. S2 in Appendix S2).

Within the set of the genera studied, the Caribbean and

the Western Palaearctic bioregions were the most diverse

with 47 and 39 fossil genera collected respectively (see

Fig. S2 in Appendix S2). The Nearctic and the Eastern

Palaearctic bioregions were less diverse with 25 and 12 fossil

genera collected respectively. The fossil records from the

Australasian, Afrotropical, Indomalayan, Malagasy or

Neotropical bioregions for the periods considered were

scarce or totally absent.

Generic overlaps

We evaluated the fraction of fossil genera of different biore-

gions that are present in the modern faunas of those same

bioregions, using genera that are present in both time peri-

ods. The palaeofauna of the West Palaearctic bioregion, for

all periods considered, shows its strongest overlap with the

modern ant fauna of the Indomalayan, Eastern Palaearctic

and Australasian bioregions (95%, 90% and 79.5% of fossil

genera are present in the modern faunas respectively). In

contrast, the fossil ant generic composition of the West

Palaearctic bioregion show limited overlap to its modern

equivalent, with only 64% of fossil genera occurring in mod-

ern faunas; a percentage similar to those observed in the

Nearctic (67%), Afrotropical (64%) or Neotropical biore-

gions (64%). Overlap values are similar between the geologi-

cal periods considered (Fig. 1).

The Caribbean ant palaeofauna has the highest overlap

with the Neotropical and Nearctic bioregions (98% and 87%

similarity of genera respectively) and to a lesser extent with

the modern Caribbean bioregion (74%). There is limited evi-

dence for overlap with other bioregions (52% to 37% simi-

larities, Fig. 1).

The fossil fauna of the Nearctic bioregion shows its stron-

gest affinity with its own modern fauna (100% overlap), the

Neotropical bioregion (91%) and to a lesser extent with the

Caribbean bioregion (83%). Other bioregions present less

affinity with the fossil fauna of the Nearctic bioregion (39–57%). Those values differ, however, for the two geological

periods considered. The Nearctic Eocene fauna presents

stronger affinities with the Eastern Palaearctic, the Indoma-

layan and the Western Palaearctic bioregions (80% overlap

of generic composition for all; Fig. 1). In contrast, the over-

lap with the Caribbean bioregion is less pronounced (60%;

Fig. 1). It is important to note that the number of fossil gen-

era known from this period is limited (n = 11). The Miocene

fauna is comparable to the overall pattern described above

with, however, lower similarity values with the Eastern and

Western Palaearctic bioregions and the Indomalayan biore-

gion, but with higher similarity value with the Caribbean

bioregion.

The fossil fauna of the East Palaearctic is most aligned

with the modern ant fauna of the Indomalayan bioregion

(Fig. 1). The affinity of the fossil and modern ant fauna of

the Eastern Palaearctic bioregion present strong affinities,

with 83% of the genera known from the fossil records still

present in this bioregion. The overlap of the East Palaearctic

fossil fauna with the modern Nearctic bioregion is also

noticeable (67%). However, the highest affinity between

those two bioregions is found between the Eocene Nearctic

palaeofauna and the modern East Palaearctic fauna (80%).

Taphonomic bias of generic records

From 74 extant genera known also to have fossil species, our

results did not support the hypothesis that arboreal genera

were overrepresented in the fossil record (Table 1, n = 96,

v2 = 0.53, P = 0.77), even when considering only the genera

known from the worker caste (n = 66, v2 = 1.09, P = 0.58).

Those results may be limited as many genera could not be

associated to a strict category (arboreal, epigaeic or hypo-

gaeic) as several of their extant species are known to occupy

more than one stratum. The multinomial logistic regression

analysis (Table 2) supports an epigaeic origin of the species

richness across subfamilies when all fossil deposits sites are

considered. This result is, however, slightly different when

fossil deposits are considered individually. While the Baltic

amber and Bitterfeld amber deposits relate more strongly to

an epigaeic or epigaeic + hypogaeic origin, the Rovno and

Scandinavian amber deposits appear biased towards an arbo-

real community composition origin.

Community structure comparison

The clustering analysis shows that the different modern com-

munities present a specific biogeographical signature that is

translated in the relative presence and species richness of the

different ant subfamilies (Fig. 2). The fossil assemblages con-

sidered also strongly clumped together indicating important

affinities in their species composition at the subfamily level

relative to modern assemblages. The fossil assemblages show

particular affinities with modern communities of the


5


Indomalayan, Australasian, and Neotropical bioregions

(Fig. 2). The West Palaearctic modern ant communities are

distinct from their fossil equivalent, a pattern also observed

with modern Nearctic and East Palaearctic communities

(Fig. 2).

The multinomial logistic regression analysis (Table 2)

shows several consistent features among all fossil deposits

based on species richness across subfamilies. The structure of

fossils deposit communities is generally most similar to mod-

ern communities in the Indomalayan and Australasian biore-

gions. The Rovno amber and Scandinavian amber deposits

also show some similarities with the modern assemblages in

the Oceania bioregion. For all fossil deposits considered or

individually, the model predicts a primary forest origin.

DISCUSSION

The aggregated fossil record reveals the dynamic nature of

macroecological patterns in ant biodiversity during the

Cenozoic. The commonness of large-scale extinctions on the

generic level is one of the most noticeable features of the

data and analyses. For example, the generic extinction rates

of the Caribbean and Western Palaearctic bioregions follow-

ing the Miocene period are at least 26% and 36% respec-

tively. In addition, from 39 genera known from the

European deposits and with modern representatives some-

where in the world, 14 (or 36%) have gone extinct from this

bioregion or remain only in the southern portion of this

bioregion.

These numbers would undoubtedly increase significantly

with the inclusion of genera only known from the fossil

record, which were excluded in our analysis. For example,

0.74

0.85

0.60

0.42

0.40

0.54

0.52

0.87

1.00

1.00

0.67

0.65

0.69

0.71

0.98

0.95

0.80

0.67

0.60

0.65

0.65

0.43

0.45

0.30

0.33

0.50

0.62

0.58

0.39

0.45

0.30

0.42

0.45

0.50

0.45

0.37

0.50

0.80

0.67

0.65

0.73

0.68

0.43

0.60

0.80

0.83

0.90

0.92

0.90

0.52

0.60

0.80

1.00

0.95

1.00

1.00

0.50

0.50

0.40

0.75

0.75

0.77

0.74

Carribean

Nearctic

Neotropical

Ethiopian

Malagasy

West Palaearctic

Indo-Malayan

East Palaearctic

Australian

Carribean Miocene

Nearctic Miocene

East Palaearctic Miocene

West Palaearctic Miocene

Nearctic Eocene

West Palaearctic Oligocene

West Palaearctic Eocene

Modern Ant Faunas

Fossil Ant Faunas

Figure 1 Generic similarity between the ant

palaeofauna known from the differentgeological periods and the modern ant

fauna within each bioregion. Lighter coloursindicate higher similarities. Values can range

from 0 (no genera in common) to 1 (allgenera known are shared).

Table 1 Number of ant genera known in the fossil record in

function of the known caste (worker or sexual) as fossil and themicrohabitat they presumably occupied (based on current

ecology of each genus).

Number of

Genera

Known from

worker caste

Known only

from sexual

caste

No data

available

Arboreal 14 12 1 1

Epigaeic 18 14 1 3

Hypogaeic 17 9 5 3

Epigaeic and

arboreal

20 15 2 3

Epigaeic and

hypogaeic

1 0 1 0

All strata 4 3 1 0

Table 2 Results of the multinomial logistic regression analysis

presenting the specific affinities for all ant fossil deposits andeach of the four ant fossil deposits studied with the modern ant

composition in species richness across subfamilies in function oftheir origin (bioregion), habitat and strata of collection.

Likelihood

value Bioregion Habitat Strata

All fossils �152.906 Indomalaya Primary forest E

�153.638 Australasia Primary forest E

�166.142 Australasia Primary forest E+H

Baltic

Amber

�82.120 Indomalaya Primary forest E



Bitterfeld

Amber



�60.256 Indomalaya Primary forest E

Rovno

Amber

�17.409 Oceania Primary forest A

�17.532 Australasia Primary forest A

�18.951 Indomalaya Primary forest A

Scandinavian

Amber

�10.626 Oceania Primary forest A

�11.367 Indomalaya Primary forest A

�12.421 Australasia Primary forest A

A, arboreal; E, epigaeic; H, hypogaeic.


6


extinct genera of the subfamily Aneuretinae are present in

European Tertiary fossil deposits but the subfamily is cur-

rently restricted to Sri Lanka (Dlussky, 2012). Similarly,

extinct ant genera of the subfamily Myrmeciinae are present

in Nearctic and European fossil faunas (Archibald et al.,

2006), but the subfamily is now restricted to Australasia.

Regional extinction also occurred even within the most ubiq-

uitous ant genus, Camponotus (Gu�enard et al., 2010), known

in the fossil record from England but now extinct in the Bri-

tish Isles. This pattern of widespread, large-scale extinction

helps to explain some of the 20 extreme biogeographical dis-

junctions in ant genera observed today (Gu�enard et al.,

2010), suggesting a broader historical distribution of those

genera. For example the genus Perissomyrmex is only known

from a few localities in Central America and Southeast Asia

(Ogata & Okido, 2007).

The fossil West Palaearctic–modern Indomalayan-

Australasian affinity

Our analyses using several data sets and analytical

approaches support the hypothesis of Emery (1892), and

Wheeler (1915) that the ant fauna of Europe during the

Eocene-Miocene has affinities with the Indomalayan biore-

gion and to a lesser extent the Australasian bioregion. In

addition to the strong generic composition similarities, we

West Palaearctic East PalaearcticNearctic

OceaniaMalagasyAfrotropical

AustralianNeotropical Indo-Malayan

NMDS Axis 1

NM

DS

Axi

s 2

Figure 2 Nonmetric multidimensional scaling of variability in ant species richness composition across subfamilies in function of thedifferent bioregions of the world. On each figure, fossil deposits communities appear in red, modern communities for the specific

bioregion appear in blue and in grey the rest of the points not specific to the studied bioregion. Size of the dot is proportional to thespecies richness with bigger dots representing richer communities.


7


also demonstrate a structural similarity in species richness of

the extant subfamilies between the Cenozoic ant fauna of

Europe and the modern ant fauna of the Indomalayan and

Australasian bioregions. Without well-studied ant fossil

records from the latter bioregions, it is difficult to determine

whether these cases represent contractions of a historically

wide-ranged genera, or whether ranges have shifted over time

into new bioregions. Studies conducted on other organisms

such as plants (Manchester et al., 2009) and bees (Michez

et al., 2009), have suggested the former. One open question

is whether the similarity of West Palaearctic faunas with Aus-

tralasia is mostly due to recent arrivals of Indomalayan lin-

eages, and thus the true historical similarity is with

Indomalaya. Although it is difficult to be conclusive with the

available data, of the 39 genera that were previously present

in the West Palaearctic and are now restricted to the Indo-

malayan or Australasian bioregions, most (30) are present in

both Indomalaya and Australasia. Eight are restricted to

Indomalaya, and only one is restricted to Australasia. Further

study, including consideration of the phylogenetic positions

of different genera and the ages of colonization events, could

more rigorously address this question.

Our analyses of community structure, the distribution of

species richness across subfamilies, also support an affinity

between the Western Palaearctic palaeofauna and modern

Indomalayan and Australasian ant fauna. First, the four fossil

assemblages used in this study present strong similarities as

shown in the clustering analysis (Fig. 2). Second, fossil

assemblages show higher affinities with the modern commu-

nities observed in the Indomalayan and Australasian biore-

gions (Fig. 1). Finally, these results are also supported by the

modelling approach where fossil assemblages from the West

Palaearctic bioregion are best predicted by models represent-

ing the Indomalayan or Australasian bioregions (Table 2).

Interestingly, the affinity of the European ant palaeofauna

with the current Afrotropical bioregion is very limited. While

seemingly counterintuitive based on the modern configura-

tion of continents, this could be explained by the separation

of the African and Eurasian plates until the middle Miocene

period (McQuarrie et al., 2003).

These results, based on three approaches, support the

hypothesis that the fossil assemblages of Europe during the

Palaeogene were most similar in generic composition and

community structure with the modern ant fauna of the

Indomalayan and Australasian bioregions. These results are

also consistent with studies in other taxa, including (but not

limited to) Salticid spiders (Pr�osz�nski & _Zabka, 1980), bees

(Engel, 2001), Hemiptera (Popov et al., 2011), and plants

(Reid & Chandler, 1933; Manchester, 1999; Manchester

et al., 2009; Teodoridis et al., 2012).

The East Palaearctic–East Nearctic disjunction

Support for the East Palaearctic–East Nearctic disjunction is

common in plants and ferns (Wen, 1999), but scarcer for

insects (for fossil examples, see Archibald & Makarkin,

2006). The generic composition of all the fossil deposits from

the Holarctic bioregion show high similarity with the current

East Palaearctic or Indomalayan bioregions supporting the

hypothesis of a widespread ant assemblage in the Northern

Hemisphere possibly facilitated by intermittent land connec-

tions accessible to both cold and warm-adapted lineages (e.g.

Archibald et al., 2011). As discussed above, the overlap

between the now-extinct West Palaearctic fauna and the

current Asian ant fauna was especially important.

Similarity between the current Neotropical bioregion

and the Miocene Caribbean fauna

Our results support Wilson’s conclusion that the Dominican

palaeofauna has strongest affinities with current Neotropical

faunas. From 47 genera known from the fossil records of

Dominican amber, only one genus, Leptomyrmex, is not

encountered in the Neotropical bioregion, and six genera are

unknown from the modern Nearctic. Leptomyrmex remains

an enigmatic case. A recent molecular phylogeny of the Doli-

choderines (Ward et al., 2010) indicated that the sister group

to Leptomyrmex is Neotropical, lending credence to the idea

that a lineage colonized Australasia from the Neotropics and

gave rise to the current genus. However, Dlussky et al.

(2014) recently argued based on morphology that the

Neotropical Leptomyrmex was nested within Old World Lep-

tomyrmex. Further work is needed to better understand this

enigma. Additionally, although subgeneric patterns are not

the focus of our analysis, recent work sheds light on a

prominent example in the genus Pheidole that influenced

Baroni Urbani’s conclusions (Baroni Urbani, 1995). The

presence of highly spinescent Pheidole (e.g. P. primigenia) in

the Dominican amber has been interpreted as presence of a

lineage that is now restricted to the Old World, as all extant

spinescent Pheidole are restricted to the IndoMalayan and

Oceanian bioregions. However, recent phylogenetic analysis

of Pheidole (Sarnat & Moreau, 2011; Economo et al., 2015)

showed that spinescence has evolved multiple times in the

Old World, thus it seems likely that the spinescent forms in

Dominican Pheidole reflect convergent evolution as intercon-

tinental dispersal and subsequent extinction, and such dis-

persal is otherwise exceedingly rare in Pheidole (Economo

et al., 2015).

Taphonomic bias

Finally, our results indicate that ant fossil assemblages are

not systematically biased towards arboreal communities. This

result is supported both in terms of composition, where no

difference in the overall presence of hypogaeic-epigaeic-arbo-

real genera was retrieved, as well as in the subfamily compo-

sition in ant assemblages. Two fossils deposits, namely the

Rovno and the Scandinavian amber deposits, show modest

support for a bias towards arboreal strata. It is unclear

whether this reflects noise due to sampling effects or the

signal of real biological differences. Other comparisons


8


between the Rovno and Baltic amber arthropods assemblages

showed that those deposits represent two different fauna,

with the Rovno amber fauna probably formed within a more

xeric subtropical forest (Perkovsky, 2009).

While we cannot rule out the presence of taphonomic

biases of various kinds based on these data alone, the lack of

a strong arboreal bias in amber fossilization is somewhat

non-intuitive. However, although resin is produced on the

trunks and branches of trees, it is also produced by roots at

the ground level or underground (Mart�ınez-Delcl�os et al.,

2004), which in principle could catch soil-dwelling ants.

Some amber deposits yield a large amount of such ‘litter

amber’ (Perrichot, 2004), providing fossils of litter-living

organisms.

CONCLUSIONS

Our results highlight the dynamic nature of biogeographical

processes shaping modern ant assemblages. In particular, our

data suggest a climate-driven reorganization of the Palaearctic

ant fauna. Many ant lineages living previously in Europe when

climate was much warmer were not able to persist through

long-term cooling and glacial cycles to the present. Interest-

ingly, the warm-adapted lineages that went extinct in Europe

can be found today in Indomalaya and Australia, but not as

often in the Afrotropics, possibly reflecting differences in conti-

nental contiguity. In contrast, the Nearctic fossil genera that

have not gone extinct are all still present in the Nearctic, with

the exception of the subfamily Myrmeciinae which was in the

Nearctic and West Palaearctic during the Eocene and is now

restricted to Australasia (Archibald et al., 2006; Dlussky, 2012).

The shift seen in Europe hints at a common explanation

for the latitudinal diversity gradient; that tropical areas have

been bigger for longer, whereas temperate areas are relatively

new biomes that may have not had time to build up diverse

floras and faunas adapted to those climates (Fischer, 1960;

Wiens & Donoghue, 2004). Relevant climate changes could

involve general cooling trends in temperature and/or an

increase in seasonality in the Northern Hemisphere (Archi-

bald & Farrell, 2003; Archibald et al., 2010).

Recent advances in the systematics and biogeography of

modern ants, and the new tools available for analysing fossils

(e.g. X-ray lCT), open a new era for the understanding of

ant biogeography and evolution. In this context, we strongly

advocate for increased research in palaeoentomology. Further

work on existing deposits, and analysis of newly discovered

fossil deposits in India (Cambay amber; Rust et al., 2010),

Australia (Cape York amber; Hand et al., 2010), France (Oise

amber), and New Zealand, will no doubt enrich our knowl-

edge of ant evolution and biogeography.

ACKNOWLEDGEMENTS

We warmly thank all the contributors to the Ant Genera of

the World Project who provided data to B.G. We thank

three anonymous referees for comments on the ms. E.P.E.

and B.G. were supported by OIST and an NSF grant to

E.P.E. (NSF DEB-1145989).

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SUPPORTING INFORMATION

Additional Supporting Information may be found in the

online version of this article:

Appendix S1 Supplementary methods and list of references

used to build the Global Ant Community Database.

Appendix S2 Supplementary tables and figures.

Appendix S3 Data from the Global Ant Community Data-

base used for the global community analysis.

BIOSKETCHES

Benoit Gu�enard is an assistant professor at the University

of Hong Kong. His research focuses on ant diversity patterns

and the mechanisms that shape them at local to global

scales.

Evan P. Economo is interested in theoretical and empirical

approaches to understanding biodiversity patterns and pro-

cesses in a geographical context. He is particularly fascinated

by the ecology, evolution, and biogeography of ants.

Vincent Perrichot is a palaeontologist interested in the

evolution of insects and the paleoecology of amber forests.

He is particularly fascinated by the origin and evolution of

ants.

Author contributions: E.P.E., B.G., and V.P. conceived the

study. B.G. and V.P. collected the data, B.G. and E.P.E. anal-

ysed the data. All authors contributed to the preparation of

the manuscript.

Editor: Lawrence Heaney


11


Journal of Biogeography


Integration of global fossil and modern biodiversity data reveals dynamism and stasis in

ant macroecological patterns

Benoit Guénard, Vincent Perrichot and Evan P. Economo

Appendix S2: Supplementary tables and figures

Table S1. List of fossil deposits where ant genera that have not gone extinct have been

collected. The geological period and age of each deposit are taken from the Paleobiology

Database (https://paleobiodb.org). Bioregions are classified according to figure S1.

Deposit name Geological period

Estimated age (Ma Years)

Country / Region

Recorded number of genera

(Recent / Total)

West Palaearctic bioregion

Oise amber Early Eocene 52-55 France 9/11

Grube Messel, Darmstadt Middle Eocene 47 Germany 4/11

Eckfeld, Germany Middle Eocene 44 Germany 2/2

Baltic amber Middle Eocene 37- 42 Poland 28/51

Bournemouth, Dorset Middle Eocene 37-40 England 2/3

Scandinavian amber Late Eocene 34-42 Denmark 16/24

Celas, Gard Late Eocene 34-35 France 2/2

Isle of Wight Late Eocene 34 England 5/10

Kuclín, near Bílina Late Eocene 34-37 Czech Rep. 1/1

Rovno amber Late Eocene 34-37 Ukraine 19/26

Brunstatt, Haut-Rhin Early Oligocene 30-34 France 4/4

Kleinkems Early Oligocene 29-34 Germany 8/10

Auxillac, Auvergne Early Oligocene 28-34 France 1/1

Aix en Provence Middle Oligocene 28 France 7/7

Bitterfeld amber Late Oligocene 23 Germany 23/39

Rott, Westphalia Late Oligocene 23 Germany 3/5

Decín Early Miocene 16-23 Czech Rep. 1/1

https://paleobiodb.org

https://paleobiodb.org

Sicilian amber Early Miocene 16-23 Sicily 10/13

Radoboj Early Miocene 19 Croatia 7/15

Mokrina Early Miocene 16-18 Czech Rep. 3/3

Vishnevaya Balka Middle Miocene 11 Southern Russia 1/4

Brünn-Vösendorf Late Miocene 7.5-9 Austria 2/2

Montagne d'Andance Late Miocene 7.5 France 2/2

Joursac, Auvergne Late Miocene 5-7 France 3/3

Caucasus region Late Miocene 5.5-11 Southern Russia 1/2

Schossnitz, Silesia Late Miocene 5-11 Poland 3/3

Oeningen Late Miocene 5.3 Switzerland 6/8

Crete Miocene 5.3-7.2 Greece 1/1

Kerch, Crimea Peninsula Miocene ? Ukraine 1/1Lac Chambon, Auvergne Pliocene 1.8-5.3 France 2/2

East Palaearctic bioregion

Shanwang Early Miocene 17 Shandong (China) 11/18

Chon-Tyz mine Middle Miocene 12-16 Kyrgyzstan 3/3

Chojaburu Middle Miocene 11.6-16 Kyushu (Japan) 1/1

Indomalayan bioregionBorneo amber Miocene Borneo 1/1

Caribbean bioregion

Dominican amber Early Miocene 16-19 Hispaniola 31/33

Nearctic bioregion

Arkansas amber Middle Eocene 40-48 Arkansas 1/3

Green River formation Middle Eocene 40-48 Colorado / Wyoming

3/12

Mossy Creek, near Wellborn

Late Eocene 34-37 Texas 1/1

Florissant Late Eocene 34 Colorado 11/19

Quesnel Early Miocene? 20-23 British Columbia 3/4

Mexican amber Middle Miocene 13-20 Chiapas 11/11

Central Alaska Late Miocene 6.7 Alaska 4/4

Afrotropical bioregion

Mfwangano Island, Lake Victoria

Early Miocene 16-23 Kenya 1/1

Table S2. Number of sites sampled for ants in each habitat type within each bioregion.

Abbreviations for the bioregions studied are as follow: Afrotropical (Afr), Australasian (Aus),

East Palaearctic (Ea Pa), Malagasy (Mal), Nearctic (Nea), Neotropical (Neo), Oceania (Oce),

Indomalayan (In-Ma), West Palaearctic (We Pa). Primary forest (=Pr. Forest).

Afr Aus Ea Pa Mal Nea Neo Oce In-Ma We PaDesert 13 1 8 0 8 1 0 0 45

Grassland 15 47 40 1 75 26 4 9 210

Pr. forest 32 97 93 27 112 324 11 129 57

Shrubland 61 66 6 0 49 39 2 5 30

Total 121 211 147 28 244 390 17 143 342

Table S3. Ecological classification of ant genera and castes from which fossil specimens have

been recovered. The following abbreviations have been used to present the castes: W=

workers, G= gyne, M= Male, U= caste unknown.

Genus name Habitat Strata Caste collectedAcanthognathus Hypogaeic WAcanthostichus Hypogaeic WAcropyga Hypogaeic G + MAnochetus ALL GAnonychomyrma Epigaeic + Arboreal W + G + MAphaenogaster Epigaeic WApterostigma Epigaeic + Arboreal WAzteca Arboreal W + G + MBrachymyrmex Epigaeic + Arboreal UCamponotus Epigaeic + Arboreal W + G + MCarebara Hypogaeic W + G + MCataulacus Arboreal W + MCephalotes Arboreal WCerapachys Hypogaeic UCrematogaster Epigaeic + Arboreal G + MCylindromyrmex Arboreal GCyphomyrmex Epigaeic W + GDiscothyrea Hypogaeic W + G + MDolichoderus Arboreal W + G + MDorymyrmex Epigaeic UEurhopalothrix Hypogaeic UForelius Epigaeic UFormica Epigaeic W + G + MGesomyrmex Arboreal W + G + MGnamptogenys ALL W + G + MGracilidris Epigaeic WHypoponera Hypogaeic G + MIridomyrmex Epigaeic + Arboreal MLasius Epigaeic W + G + MLeptogenys Epigaeic + Hypogaeic GLeptomyrmex Epigaeic + Arboreal WLinepithema Epigaeic + Arboreal ULiometopum Epigaeic W + G + MMeranoplus Epigaeic UMessor Epigaeic GMonomorium Epigaeic WMycocepurus Epigaeic U

Myopopone Hypogaeic GMyrmelachista Arboreal UMyrmica Epigaeic W + G + MNeivamyrmex Hypogaeic WNesomyrmex Arboreal WNylanderia Epigaeic + Arboreal W + G + MOctostruma Hypogaeic UOdontomachus Epigaeic + Arboreal WOecophylla Arboreal W + G + MParaponera Epigaeic + Arboreal WPheidole ALL WPlagiolepis Epigaeic + Arboreal W + G + MPlatythyrea Arboreal W + GPodomyrma Arboreal WPogonomyrmex Epigaeic WPolyrhachis Epigaeic + Arboreal WPonera Hypogaeic G + MPrenolepis Epigaeic + Arboreal GPrionopelta Hypogaeic UPristomyrmex Epigaeic WProceratium Hypogaeic W + G + MPseudolasius Epigaeic WPseudomyrmex Arboreal W + G + MRhytidoponera Epigaeic WSolenopsis ALL W + G + MStenamma Hypogaeic MStigmatomma Hypogaeic W + MStrumigenys Hypogaeic W + GTapinoma Epigaeic + Arboreal W + MTechnomyrmex Arboreal WTemnothorax Epigaeic + Arboreal W + MTetramorium Epigaeic + Arboreal UTetraponera Arboreal W + G + MTrachymyrmex Epigaeic WVollenhovia Epigaeic + Arboreal WWasmannia Epigaeic + Arboreal UZatania Epigaeic + Arboreal W + M

Figure S1. Map of the bioregions used in this study. Divisions of the world into multiple

administrative and geographic regions follow the classification of Guénard et al. 2012.

Reference

Guénard, B., Weiser, M.D. & Dunn, R.R. (2012) Global models of ant diversity suggest

regions where new discoveries are most likely are under disproportionate deforestation threat.

Proceedings of the National Academy of Sciences USA, 109, 7368–7373.

West Palaearctic

East Palaearctic

Indomalayan

Australasian

Afrotropical

Malagasy

Caribbean

Nearctic

Neotropical

Figure S2. Summary of the known distribution for all genera of ants known from fossil

deposits and with current distribution. Each row for a given genus indicates its presence for a

given bioregion. If coloured then presence known for the specific period, otherwise if blank

then presence unknown. Abbreviations are as follow for the geological periods: E= Early, M=

Middle and L= Late. Bioregions are classified according to figure S1.

Journal of Biogeography


Integration of global fossil and modern biodiversity data reveals dynamism and stasis in

ant macroecological patterns

Benoit Guénard, Vincent Perrichot and Evan P. Economo

Appendix S1: Supplementary methods

Assembly of the fossil and modern biodiversity databases

Our dataset of fossil specimens was based on the list established by one of the authors

for AntWeb (Perrichot 2014). This list was assembled through a literature search for generic

mentions (Brown 1973, Wilson 1985, Lattke 1990, Baroni Urbani & De Andrade 1994, 2003,

de Andrade 1994, 1995, 1998, Baroni Urbani, 1995, Poinar et al. 1999, 2006, Baroni Urbani

& De Andrade 2003, Dlussky & Rasnitsyn 2009, Perkovsky 2009, Heterick & Shattuck 2011,

Blaimer 2012, Durán-Ruiz et al. 2013, Wappler et al. 2013, Antropov et al. 2014, Dlussky &

Putyatina 2014), and also complemented by unpublished records from the personal

observations by V.P. during his work on institutional and private collections. No deposit older

than the early Eocene is known to have fossils of modern ant genera, which limited the

number of included deposits. The modern distributions of the genera were extracted from

Guénard et al. (2010, 2012), complemented by the recent relevant taxonomic modifications

(LaPolla et al. 2010, Yoshimura & Fisher 2012), and assigned to specific bioregions.

Definition of the biogeographic regions

The Nearctic bioregion is defined here to include all the regions north of Mexico and

inclusive of Mexico. Mexico is included to this bioregion on the basis of palaeogeological

results suggesting a direct connection between North America and Mexico from the

Paleocene to the Miocene, but not to South America, as the Panama isthmus was not formed

yet (Keigwin 1978). However, to fully consider the hypothesis of a more distinct ant fauna

(genera) associated with Mexico and not with the rest of the Nearctic bioregion, we tested the

placement of southern Mexico (all the states located south or east of Oaxaca and Veracruz

states, with these later two included) within the Neotropical bioregion. As a result, only one

genus (Cylindromyrmex, known only from the Caribbean fossil fauna) was excluded from the

current Nearctic bioregion. Finally, these alternative classifications of the bioregions did not

modify substantially the final results or our conclusions.

Taxonomic classification

Our taxonomic classification followed Bolton (2013), and we took a conservative

approach by excluding doubtful records. The record of Ectatomma by Emery (1891) from

Sicilian amber was transferred to Gnamptogenys following Brown and Carpenter (1978). The

genera Mycetosoritis and Stenamma were mentioned by Brown (1973) respectively on the

base of “possible “male and “shrunken and distorted” specimens from Chiapas amber.

However, the validity of these identifications was later questioned (Baroni Urbani 1980,

Branstetter 2012) and never validated to our knowledge, thus we excluded them from our

analysis. Records of Pachycondyla specimens from Eocene amber and imprints (Dlussky

2002, Dlussky & Rasnitsyn 2009, Aria et al. 2011, Dlussky & Wedmann 2012) were not

included in our analysis as their current placement is unresolved in the recent taxonomic

changes of the genus (Schmidt & Shattuck 2014).

It should be noted that several controversies on the placement or validity of specific

fossil species or genera exist; with several deposits receiving incomplete or now outdated

treatments (e.g. Fushun deposit, Fujian, China)(LaPolla et al. 2013). The data used for the

analyses performed here reflect the current state of ant taxonomy and that future work on both

fossil and modern taxa could modify the current framework (e.g. the recent changes within

the Dorylinae: Brady et al. 2014, or Myrmicinae subfamilies: Ward et al. 2015).

Community database

The literature database was compiled through extensive searches by B.G. during

recent projects on the global distribution of Formicidae. Only publications that provided a list

of species collected were used so that we could keep the taxonomy up to date (e.g. the

Amblyoponinae, Ectatominae and Proceratiinae subfamilies were once sunk within the

Ponerinae subfamily).

From the 1060 studies with data for 4376 sites, we excluded sites for which only

baiting methods were used, which can result in a bias toward more dominant and generalist

species (Bestelmeyer et al. 2000). We selected habitats representative of undisturbed or with

limited natural disturbance and from which habitat could be categorised as one of the

following category: primary forest, shrubland, grassland, or desert. Finally, we also excluded

communities with a species richness lower than ten from the NMDS analysis, as their

proportions among subfamilies will be noisy due to sampling effects.

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