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Systematic Entomology (2015), 40, 61–81 DOI: 10.1111/syen.

12090

The evolution of myrmicine ants: phylogeny


and biogeography of a hyperdiverse ant clade
(Hymenoptera: Formicidae)
P H I L I P S . W A R D 1, S E Á N G . B R A D Y 2, B R I A N L . F I S H E R 3
and T E D R . S C H U L T Z 2
1
Department of Entomology and Nematology, University of California, Davis, CA, U.S.A., 2 Department of Entomology, National
Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A. and 3 Department of Entomology, California Academy
of Sciences, San Francisco, CA, U.S.A.

Abstract. This study investigates the evolutionary history of a hyperdiverse clade, the
ant subfamily Myrmicinae (Hymenoptera: Formicidae), based on analyses of a data
matrix comprising 251 species and 11 nuclear gene fragments. Under both maximum
likelihood and Bayesian methods of inference, we recover a robust phylogeny that
reveals six major clades of Myrmicinae, here treated as newly defined tribes and occur-
ring as a pectinate series: Myrmicini, Pogonomyrmecini trib.n., Stenammini, Solenop-
sidini, Attini and Crematogastrini. Because we condense the former 25 myrmicine tribes
into a new six-tribe scheme, membership in some tribes is now notably different, espe-
cially regarding Attini. We demonstrate that the monotypic genus Ankylomyrma is nei-
ther in the Myrmicinae nor even a member of the more inclusive formicoid clade – rather
it is a poneroid ant, sister to the genus Tatuidris (Agroecomyrmecinae). Several
species-rich myrmicine genera are shown to be nonmonophyletic, including Pogono-
myrmex, Aphaenogaster, Messor, Monomorium, Pheidole, Temnothorax and Tetramor-
ium. We propose a number of generic synonymies to partially alleviate these problems
(senior synonym listed first): Pheidole = Anisopheidole syn.n. = Machomyrma syn.n.;
Temnothorax = Chalepoxenus syn.n. = Myrmoxenus syn.n. = Protomognathus syn.n.;
Tetramorium = Rhoptromyrmex syn.n. = Anergates syn.n. = Teleutomyrmex syn.n. The
genus Veromessor stat.r. is resurrected for the New World species previously placed
in Messor; Syllophopsis stat.r. is resurrected from synonymy under Monomorium to
contain the species in the hildebrandti group; Trichomyrmex stat.r. is resurrected from
synonymy under Monomorium to contain the species in the scabriceps- and destruc-
tor-groups; and the monotypic genus Epelysidris stat.r. is reinstated for Monomorium
brocha. Bayesian divergence dating indicates that the crown group Myrmicinae origi-
nated about 98.6 Ma (95% highest probability density 87.9–109.6 Ma) but the six major
clades are considerably younger, with age estimates ranging from 52.3 to 71.1 Ma.
Although these and other suprageneric taxa arose mostly in the middle Eocene or earlier,
a number of prominent, species-rich genera, such as Pheidole, Cephalotes, Strumigenys,
Crematogaster and Tetramorium, have estimated crown group origins in the late Eocene
or Oligocene. Most myrmicine species diversity resides in the two sister clades, Attini
and Crematogastrini, which are estimated to have originated and diversified extensively
in the Neotropics and Paleotropics, respectively. The newly circumscribed Myrmicini is
Holarctic in distribution, and ancestral range estimation suggests a Nearctic origin. The
Pogonomyrmecini and Solenopsidini are reconstructed as being Neotropical in origin,

Correspondence: Philip S. Ward, Department of Entomology and Nematology, University of California, Davis, CA 95616, U.S.A. E-mail:
psward@ucdavis.edu

© 2014 The Royal Entomological Society 61


62 P. S. Ward et al.

but they have subsequently colonized the Nearctic region (Pogonomyrmecini) and many
parts of the Old World as well as the Nearctic region (Solenopsidini), respectively. The
Stenammini have flourished primarily in the northern hemisphere, and are most likely
of Nearctic origin, but selected lineages have dispersed to the northern Neotropics and
the Paleotropics. Thus the evolutionary history of the Myrmicinae has played out on a
global stage over the last 100 Ma, with no single region being the principal generator of
species diversity.

This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:


zoobank.org:pub: BB6829C4-DA79-45FE-979E-9749E237590E.

Introduction generate a robust picture of the evolutionary history of these


remarkable ants. These results also provide the basis for a
Ants are one of evolution’s great success stories, the most revised higher classification of the subfamily.
species-rich and ecologically dominant of all eusocial insects
(Hölldobler & Wilson, 1990; Lach et al., 2009), yet this success
is far from evenly distributed among the major lineages of the Material and methods
group. There are 16 extant subfamilies of ants (Bolton, 2014;
Brady et al., 2014), but four of these – Dolichoderinae, Formic- Taxon selection
inae, Myrmicinae and Ponerinae – account for almost 90% of
all known species (Bolton, 2014). Among these ‘big four’ the We assembled a set of 234 species representing all 25 tribes of
most biologically diverse and prolific is the subfamily Myrmic- Myrmicinae and 128 of the 145 extant genera (taxonomy follows
inae. With approximately 6475 described species – about half Bolton, 2014) (Table S1). One species could not be identified to
of all named ants – and many others awaiting discovery and genus. For genera with widespread distributions we attempted
description, the myrmicines are a hyperdiverse clade inhabiting to sequence a species from each of the major biogeographic
most of the terrestrial surface of the Earth and encompassing a regions in which the genus occurs, to facilitate ancestral range
wide range of lifestyles including generalist and specialist preda- estimation. We also sampled more intensively within several
tors, scavengers, omnivores, granivores and herbivores (Kugler, genera whose monophyly appeared to be questionable, includ-
1979; Brown, 2000). ing Aphaenogaster, Monomorium, Pheidole, Solenopsis, Stru-
Myrmicine ants have been subjects of intense investigation migenys and Tetramorium. We included 17 outgroup taxa; these
into colony organization, communication, demography and were drawn mostly from the other subfamilies of ants, but we
ecology (Davidson, 1977; Brian, 1983; Tschinkel, 2006). They also chose an aculeate hymenopteran wasp, Apterogyna (Bra-
contain a diverse array of socially parasitic species (Beibl dynobaenidae), as the most distant outgroup to root the tree.
et al., 2005; Buschinger, 2009) and they furnish numerous
examples of symbioses with other organisms (Davidson &
McKey, 1993; Fiedler, 2006; Russell et al., 2009), including Sequencing and sequence annotation
the iconic mutualism between fungus-growing ants and their
cultivars (Mueller et al., 2001; Hölldobler & Wilson, 2011). We sequenced fragments of 11 nuclear genes, including
Genomic studies have begun to probe the genetic architecture two ribosomal genes, 18S rDNA and 28S rDNA, and nine
of prominent myrmicines in the genera Solenopsis (Wurm protein-coding genes: abdominal-A (Abd-A), elongation factor
et al., 2011), Pogonomyrmex (Smith et al., 2011), Acromyrmex 1-alpha F1 copy (EF1aF1), elongation factor 1-alpha F2 copy
(Nygaard et al., 2012) and Atta (Suen et al., 2011). (EF1aF2), long wavelength rhodopsin (LW Rh), arginine kinase
Despite the attention that they have garnered, myrmicines (ArgK), topoisomerase 1 (Top1), ultrabithorax (Ubx), wingless
present a number of taxonomic challenges. There are many dif- (Wg) and rudimentary (CAD). Protocols for extraction, ampli-
ficult species complexes (Seifert, 2003; Schlick-Steiner et al., fication and sequencing follow Ward & Downie (2005), Brady
2006; Blaimer, 2012a), as well as multiple genera and tribes et al. (2006), Ward et al. (2010), and Ward & Sumnicht (2012).
whose monophyly is doubtful (Brady et al., 2006; Ward, 2011). Sequences were collated in Sequencher v4.10.1 (Gene
The members of this subfamily have received only modest cov- Codes Corporation, Ann Arbor, MI, U.S.A.), aligned with
erage in molecular phylogenetic studies of ants as a whole, and Clustal X v1.81 (Thompson et al., 1997), and manually edited
results have been inconsistent between studies (Brady et al., and concatenated with MacClade v4.08 (Maddison & Mad-
2006; Moreau et al., 2006). Here we aim to address this prob- dison, 2005). Alignment was straightforward for exons of
lem by employing 11 nuclear genes and comprehensive taxon protein-coding genes and for most 18S rDNA sequence. We
sampling across the entire subfamily. We combine phylogenetic excluded ambiguously aligned regions of 18S and 28S, introns
inference with divergence dating and biogeographic analysis to of the protein-coding genes, and autapomorphic indels in exons

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


Phylogeny and evolution of myrmicine ants 63

Table 1. Sequence characteristics of the 251-taxon data matrix (233 Table 2. The 21 partitions and models identified by PartitionFinder
ingroup taxa). and used in the maximum likelihood and Bayesian analyses of the
concatenated, unmodified 8987-bp dataset.
Gene No. of sites VS PS VS, ingroup PS, ingroup
Partition Blocks Model
Abd-A 600 262 221 246 206
EF1aF2 517 206 193 204 190 p1 Abd-A pos1, Ubx pos1 SYM+I+G
LW Rh 458 279 250 269 239 p2 Abd-A pos2, Ubx pos2 TVM+I+G
ArgK 673 332 297 309 285 p3 Abd-A pos3 TIM+I+G
Top1 880 477 424 464 419 p4 EF1aF2 pos3, Top1 pos3 TrNef+I+G
Ubx 627 301 254 286 232 p5 EF1aF2 pos1 GTR+I+G
Wg 409 251 218 224 197 p6 ArgK pos2, EF1aF2 pos2 TIMef+I+G
EF1aF1 359 141 129 138 128 p7 LW Rh pos3, Ubx pos3 TrNef+I+G
18S 1850 224 106 184 85 p8 LW Rh pos1 TVM+I+G
28S 1653 480 271 425 223 p9 LW Rh pos2 GTR+I+G
CAD 961 605 532 592 519 p10 ArgK pos3 TrNef+I+G
All genes 8987 3558 2895 3341 2723 p11 ArgK pos1 SYM+I+G
p12 Top1 pos1 SYM+I+G
VS = variable sites; PS = parsimony-informative sites. p13 CAD pos2, Top1 pos2 GTR+I+G
p14 Wg pos3 GTR+I+G
of Abd-A, ArgK, CAD and Wg. The resulting matrix that was p15 Wg pos1, Wg pos2 TrNef+I+G
used for all subsequent analyses contains 8987 bp (3558 variable p16 EF1aF1 pos2 JC
p17 EF1aF1 pos3 TIM+G
sites, 2895 parsimony-informative sites). This matrix (251 taxa
p18 18S, EF1aF1 pos1 TrNef+I+G
by 11 genes) contains no missing gene fragments. Of the 2761 p19 28S GTR+I+G
fragments, 499 were previously published (Ward & Downie, p20 CAD pos3 GTR+I+G
2005; Brady et al., 2006; Ward, 2007a; Branstetter, 2009; p21 CAD pos1 K80+I+G
Lucky & Sarnat, 2010; Ward et al., 2010; Brady et al., 2014);
the remaining 2262 sequences were generated for this study
(GenBank accession numbers KJ859686-KJ861947). Sequence (Table 2), which were then employed in intensive Bayesian
characteristics of each gene are summarized in Table 1. The and maximum-likelihood analyses. In cases where the model
aligned data matrix has been deposited in TreeBase (study specified by PartitionFinder was not implemented in MrBayes,
accession 15764). the next-most-complex available model was employed.
The choice of partitions for separate, single-gene Bayesian
Phylogenetic analyses analyses of each of the nuclear genes followed an iterative
scheme similar to that described above for the full concatenated
Data partitions dataset. An initial Bayesian analysis was conducted employing
We employed an iterative strategy to arrive at the partitioning partitions based on variability of codon sites. The resulting
scheme for analysing the 11-gene, concatenated dataset. In a tree was submitted to PartitionFinder using ‘models = mrbayes,’
first round of analyses we partitioned the data into 25 partitions ‘model-Selection = BIC,’ and ‘search = all.’ If PartitionFinder
based on the variability of codon-position sites within each gene. identified a partitioning scheme different from the one employed
Four genes (Abd-A, EF1aF1, EF1aF2, Ubx) were each divided in the first analysis, a second Bayesian analysis was conducted
into two partitions consisting of (i) codon positions 1 and 2 and employing the PartitionFinder scheme (Table S3).
(ii) codon position 3, resulting in eight partitions; five genes
(LW Rh, ArgK, Top1, Wg, CAD) were assigned site-specific Treatments addressing base frequency heterogeneity
models in which each codon position formed a separate parti- For each of the three codon positions within each protein-coding
tion, resulting in 15 partitions; and two nonprotein-coding genes gene, as well as for 18S and 28S, each treated as single
(18S, 28S) were each assigned a single partition, resulting in two partitions, we evaluated the homogeneity of base frequencies
partitions. The choice of nucleotide substitution model for each across taxa using PAUP* 4.0a128 (Swofford, 2002), which
partition (Table S2) was determined using the Akaike Informa- indicated that nine partitions – the third positions of each
tion Criterion (AIC) (Posada & Buckley, 2004) as implemented protein-coding gene – contained significantly heterogeneous
in JModelTest v2.0.2 (Posada, 2008). The tree resulting from base frequencies (Table S4). In order to examine the effects
a Bayesian analysis (MrBayes v3.2.1; Ronquist et al., 2012) of base frequency heterogeneity on phylogenetic results, we
of these data employing these partitions and models was then subjected each dataset to a number of treatments, including:
submitted to PartitionFinder v1.0.1 (Lanfear et al., 2012) along (i) ‘partial RY coding,’ in which, for each gene, the subset of
with 29 predefined blocks, encompassing the first, second and taxa (differing for each gene) with heterogeneous third positions
third codon positions for each of the nine protein-coding genes deviating significantly from the observed averages were coded
and the two nonprotein-coding genes (18S, 28S). Under the as RY and in which the remainder of third positions were coded
Bayesian Information Criterion (BIC), the ‘greedy’ algorithm, normally (ACTG); (ii) ‘complete RY coding’ in which all third
and ‘models = all,’ PartitionFinder identified 21 partitions positions were coded as RY; (iii) exclusion of a minimum subset

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


64 P. S. Ward et al.

of 324 characters (not necessarily third positions) deviating and a minimally defined topological constraint. We included
significantly from observed base-frequency averages based on Ankylomyrma in this constraint because of a novel result (see
the results of a likelihood-based method for filtering potential below) placing it close to Tatuidris.
nonstationary sites, to be described elsewhere (D. Swofford,
personal communication); and (iv) exclusion of 843 characters Bayesian analyses
(not necessarily third positions), the 10% deviating the most
We conducted Bayesian analyses using MrBayes v3.2.1 and
from observed base-frequency averages, also based on the same
3.2.2 (Ronquist et al., 2012) with nucmodel = 4by4, nruns = 2,
filtering method. All analyses were carried out in MrBayes
nchains = 8, samplefreq = 1000, and either 20 million or 40 mil-
v3.2.1 and v3.2.2 (Ronquist et al., 2012). For treatment i, ‘partial
lion generations and burn-ins of 2 and 4 million generations,
RY,’ decisions about whether to code a particular third-position
respectively; single-gene analyses consisted of 10 million gen-
site as ACTG or RY were made based on analyses of output
erations and burn-ins of 1 million generations. For partitioned
from PAUP* 4.0a128 (Swofford, 2002) (Table S4). Characters
analyses all parameters, including branch-length rate multipli-
were recoded as RY in Mesquite v2.73 (Maddison & Maddison,
ers, were unlinked across partitions; the only exceptions were
2010). Partitioning schemes for each of these treatments resulted
branch lengths and topology, which were linked. All analy-
from the same iterative strategy described in the Data partitions
ses were carried out using parallel processing (one chain per
section above.
CPU) on the Smithsonian NMNH LAB Topaz computing clus-
ter (Apple computers with Intel processors). To address known
Treatments addressing possible ‘wildcard’ taxa problems with branch-length estimation in MrBayes (Marshall
In order to explore the reasons for poor support in one region of et al., 2006; Brown et al., 2010; Marshall, 2010; Ward et al.,
the tree (Crematogastrini), particularly with respect to a number 2010), we set brlenspr = unconstrained:Exp(100). Burn-in, con-
of unusually long-branched taxa in that region, we conducted vergence and stationarity were assessed using Tracer v1.5 (Ram-
analyses in which subsets of 11 taxa suspected of behaving as baut & Drummond, 2009), by examining PSRF values and .stat
‘wildcards’ were excluded (Nixon & Wheeler, 1992; Kearney, output files in MrBayes, and by using Bayes factor compar-
2002). Unlike the most common type of wildcard taxa, how- isons of harmonic-mean marginal likelihoods of pairs of runs
ever, none of the taxa in our analyses contained missing data. with standard error estimated using 1000 bootstrap pseudorepli-
For this purpose we created and analysed two datasets consist- cates in Tracer 1.5 (Rambaut & Drummond, 2009), which
ing of: (i) 248 taxa, excluding Acanthomyrmex ferox, Mayriella employs the weighted likelihood bootstrap estimator of New-
ebbei and Xenomyrmex floridanus; and (ii) 243 taxa, exclud- ton & Raftery (1994) as modified by Suchard et al. (2001). For
ing Cardiocondyla mauritanica, Cardiocondyla MY01, Cardio- the full concatenated dataset, stationarity was regularly obtained
condyla thoracica, Liomyrmex gestroi, Melissotarsus insularis, within the first few million generations, even for the various
Metapone madagascarica, Metapone PG01 and Ocymyrmex cf. base-frequency-heterogeneity treatments; single-gene analyses
fortior. achieved stationarity within the first few hundred thousand gen-
erations. The results reported here are based on the combined
Single-gene analyses post-burn-in data from both runs.
In order to assess the possibility of conflicting gene trees and
to compare gene-specific support for clades, we conducted Maximum likelihood
iterative analyses of each of the 11 genes as described above, ML analyses were carried out in GARLI v2.0 (Zwickl, 2006)
using partitions and models identified by PartitionFinder v1.0.1 using parallel processing on the Smithsonian Hydra supercom-
(Lanfear et al., 2012), summarized in Table S3. All single-gene puter (Linux-based with AMD processors). Unconstrained and
analyses were conducted under Bayesian criteria as described constrained ML best-tree analyses consisted of 200 and 100
below and consisted of 10 million generations with a burn-in of replicate searches, respectively, and deviated from the default
1 million generations. settings as follows: topoweight = 0.01; brlenweight = 0.002.
ML bootstrap analyses consisted of 1000 pseudoreplicates
Constraint analyses and deviated from default settings as follows: genthreshfor-
One surprising result of several prior studies (Brady et al., 2006; topoterm = 5000; scorethreshforterm = 0.10; startoptprec = 0.5;
Moreau et al., 2006; Rabeling et al., 2008), that the species minoptprec = 0.01; numberofprecreductions = 1; treerejection-
Tatuidris tatusia is a poneroid and as such only very distantly threshold = 20.0; topoweight = 0.01; brlenweight = 0.002. In all
related to the Myrmicinae, has failed to significantly influence analyses the value for modweight was calculated as 0.0005 ×
the posterior probabilities of some myrmecologists (e.g. Baroni (#subsets + 1) (D. Zwickl, personal communication).
Urbani & De Andrade, 2007; Keller, 2011). For this reason,
we conducted a constraint analysis to explicitly quantify the
difference in likelihoods between the unconstrained outcome, Divergence dating
in which (Tatuidris tatusia + Ankylomyrma coronacantha) are
poneroids, and the outcome in which they are constrained to be We inferred divergence dates with beast v1.7.5 (Drum-
members of the Myrmicinae. The latter analysis was carried out mond et al., 2012) under a parallel configuration using BEA-
in GARLI v2.0 as described below, using 100 search repetitions GLE v1.0 on the aforementioned NMNH LAB Topaz cluster.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


Phylogeny and evolution of myrmicine ants 65

We employed an uncorrelated lognormal relaxed clock model by substantially reducing the number of nonmonophyletic
(Drummond et al., 2006). Data partitioning and nucleotide sub- genera, and by ensuring that all tribes are monophyletic.
stitution models were determined using PartitionFinder v1.0.1 In synonymizing or resurrecting genera we list all new
(Lanfear et al., 2012) in the same manner as for the MrBayes or revived combinations for the affected species, and pro-
analyses described above, except that only models available vide new names (if necessary) where secondary homonymy
in beast were evaluated. Substitution models were unlinked is created.
and clock and tree models were linked among partitions.
The tree prior was a Yule process with a ML ultrametric
starting tree. We provided a priori age distributions for 27 Phylogenetic results
nodes from the fossil record (20 in Myrmicinae, 7 in the
outgroups), based on stem-group membership (Table 3). We The broad features of phylogenetic relationships within the
conducted beast runs each for 100 million generations with Myrmicinae are stable and strongly supported across multiple
a burn-in of 25 million generations, saving trees and other methods of phylogenetic inference. Alternate Bayesian anal-
parameter values every 10 000 generations. Stationarity and yses, including RY coding for third positions, exclusion of
burn-in were determined by observing high ESS values and the some base frequency heterogeneous positions, and exclusion of
consistency of likelihood values and other results among inde- ‘wildcard’ taxa, had little or no effect on overall tree topol-
pendent runs. We combined results from four independent runs, ogy or on support for the major clades (Fig. 1, Figure S1;
obtaining a total of 300 million post burn-in generations, and Tables 5, S6). Single-gene trees were less well resolved, espe-
visualized the resulting topology and divergence times using cially those with relatively few phylogenetically informative
FigTree v1.4.0. sites, but they did not yield strongly supported clades that
conflicted with the results from concatenated analyses (Figure
S2). In the discussion below the support values that are cited
Biogeographic inference for any given node are (i) the Bayesian posterior probability
(PP) based on the concatenated, partitioned MrBayes analy-
For the purpose of inferring the distributional history of sis, and (ii) the ML bootstrap percentage (BS) based on the
Myrmicinae we adopted a coarse classification of six clas- concatenated, partitioned GARLI analysis with 1000 bootstrap
sic biogeographic areas: the Neotropical, Nearctic, Palearc- pseudoreplicates.
tic, Afrotropical, Indomalayan and Australasian regions (Cox, We find that the ectaheteromorph clade, represented in our
2001), with Wallace’s line (Wallace, 1876) as the bound- dataset by Acanthoponera and Rhytidoponera, is the sister
ary between the last two. To estimate ancestral areas we group of Myrmicinae, to the exclusion of the Formicinae.
used the likelihood-based program lagrange v20130526 (Ree This relationship has weak to moderate support (PP 0.91 BS
et al., 2005; Ree & Smith, 2008). This program implements a 64) and mirrors the findings of a recent five-gene study of
dispersal-extinction-cladogenesis (DEC) model and evaluates Formicidae as a whole (Moreau & Bell, 2013). The subfamily
the likelihood of different range inheritance scenarios at the Myrmicinae is consistently recovered as monophyletic, with one
internal nodes of an ultrametric tree. The input for this analy- caveat: the enigmatic ant genus Ankylomyrma (Bolton, 1973),
sis was (i) the post-burn-in consensus tree produced by beast hitherto placed in its own tribe within the Myrmicinae, is not
(above), with branch lengths in units of time and outgroups a member of this subfamily, nor even of the more inclusive
removed, and (ii) a matrix of distribution ranges for all termi- formicoid clade (Ward, 2007b). Rather, Ankylomyrma is sister
nal taxa (Table S5). Maximum ancestral range size was set at to the genus Tatuidris (Agroecomyrmecinae), a poneroid ant
three areas, because the overwhelming majority of contempo- that was also previously placed erroneously in the Myrmicinae
rary species occupies only one biogeographic area, and none is (Bolton, 2003). The Ankylomyrma/Tatuidris clade has very
found in more than three areas (Table S5). We also incorporated strong support (PP 1.00 BS 100) and the results of the constraint
temporal and spatial constraints, such that dispersal was con- analysis reveal that the most likely tree in which Tatuidris and
fined to adjacent areas and, in accordance with changing paleo- Ankylomyrma are constrained to be members of the Myrmicinae
geography (Scotese, 2010), different rates were used for the time has a likelihood that is lower than that of the unconstrained
periods 0–50 and 50–100 Ma (Table 4). topology by a difference of 358.66 natural-log likelihood units,
translating to a highly significant Bayes Factor of 717.32
(Nylander et al., 2004). This is another striking case where
Taxonomic changes worker ant morphology led to a mistaken inference about true
affinities (Brady et al., 2006; Moreau et al., 2006; Ward, 2011;
We propose a number of changes to the higher-level taxon- Brady et al., 2014).
omy of the Myrmicinae that are impelled by the phylogenetic With Ankylomyrma excluded, the myrmicines are a robust
findings. The ultimate goal is a phylogenetic classification in monophyletic group (PP 1.00 BS 98) consisting of six major
which all higher taxa of a given rank (in this case, genus or clades that emerge with strong support across all modes of
tribe) are monophyletic and hence mutually exclusive (Ward, analysis (Table 5), and in a consistent relationship to one another.
2011). This goal cannot yet be fully realized, but the classi- These six clades form a pectinate series, here discussed in
fication proposed here goes some distance towards this ideal sequence.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


66 P. S. Ward et al.

Table 3. List of a priori age distributions applied to 27 fossil calibration points for beast divergence dating analysis.

95%
Taxon (stem-group) Offset Median quartile Notes

Acanthognathus 15 40 73 Acanthognathus poinari in Dominican amber (Baroni Urbani & De Andrade, 1994).
Upper bound is approximate estimated age of ‘core Myrmicinae’ (Brady et al., 2006).
Aphaenogaster 42 52 73 Three described Aphaenogaster species from Baltic amber are similar in form, size and
occidentalis + Goniomma sculpture to the extant species A. subterranea (Wheeler, 1915). Calibration follows
Branstetter (2012).
Apterostigma 15 45 73 Two species of Apterostigma in Dominican amber (Schultz, 2007). Median age from
Schultz & Brady (2008).
Carebara 42 52 73 Carebara antiqua in Baltic amber (Mayr, 1868; Wheeler, 1915). Upper bound is
approximate estimated age of ‘core Myrmicinae’ (Brady et al., 2006).
Cataulacus 30 45 73 Two species of Cataulacus described from Sicilian amber (Emery, 1891).
Cephalotes atratus 15 40 73 Twelve described species of Cephalotes in Dominican amber, assignable to extant species
groups within the genus (De Andrade & Baroni Urbani, 1999). Cephalotes atratus
group is sister to all other Cephalotes (Price et al., 2014).
Crematogaster acuta 15 28 35 Crematogaster crinosa-group in Dominican amber (Blaimer, 2012b). The calibrated node
represents crown Crematogaster (Orthocrema). Median age from Blaimer (2012b).
Myrmica 42 60 85 Myrmica species from Baltic and Saxonian ambers (Dlussky & Rasnitsyn, 2009), not
definitely assignable to modern species groups, although similar to the plesiotypic
ritae-group (Radchenko et al., 2007).
Nesomyrmex 15 40 73 Two described species of Nesomyrmex from Dominican amber, similar to N. wilda and
(wilda + echatinodis) N. echinatinodis, respectively (De Andrade et al., 1999), here used conservatively to
calibrate the node representing the most recent common ancestor (MRCA) of the New
World and Old World species.
Octostruma 15 40 73 Undescribed Octostruma in Dominican amber (Wilson, 1988).
Pheidole 34 45 73 Pheidole from Florissant Formation (Carpenter, 1930).
Pogonomyrmex (striatin- 34 50 73 Pogonomyrmex fossilis (worker) from the Florissant Formation (Carpenter, 1930) is
odis + subdentatus) presumably part of a Nearctic subclade of Pogonomyrmex, here used conservatively to
calibrate the MRCA of Pogonomyrmex (excluding the southern Neotropical
P. angustus) and Hylomyrma.
Pristomyrmex 42 52 73 Pristomyrmex in late Eocene Danish amber (Dlussky & Radchenko, 2011).
Solenopsis (fugax + xyloni) 15 40 73 Undescribed Solenopsis in Dominican amber (Wilson, 1988) ascribed to ‘Solenopsis
(Diplorhoptrum)’ and ‘Solenopsis (Solenopsis)’, i.e. thief ants and fire ants,
respectively. Here employed conservatively to calibrate the node representing the
MRCA of Solenopsis papuana and S. xyloni (the latter is a Solenopsis in the fire ant
group).
Stenamma 42 52 73 Stenamma berendti (male) in Baltic amber, not assignable to species groups within the
genus (Branstetter, 2012). Calibration follows Branstetter (2012).
Strumigenys ambatrix 15 40 73 Four described species of Strumigenys from Dominican amber (Baroni Urbani & De
Andrade, 1994, 2007), with two of these (S. electrina and S. poinari) evidently nested
well within the genus, related to Nearctic taxa formerly placed in Pyramica. These are
used conservatively to calibrate the node representing crown Strumigenys.
Temnothorax 42 52 73 Six described species of Temnothorax in Baltic amber (Dlussky & Rasnitsyn, 2009).
Temnothorax poeyi 15 40 73 Temnothorax of the ‘Macromischa’ type from Dominican amber (De Andrade, 1992).
Tetramorium 42 52 73 Three undescribed species of Tetramorium in Baltic amber (Dlussky & Rasnitsyn, 2009).
Trachymyrmex 15 20 73 Trachymyrmex in Dominican amber (Baroni Urbani, 1980). Median age from Schultz &
Brady (2008).
Outgroups
Amblyoponinae 42 70 90 Stigmatomma groehni in Baltic amber (Dlussky, 2009). The divergence between two
subclades of Amblyoponinae (one containing Amblyopone, and the other
Stigmatomma) is evidently quite old, based on high levels of genetic divergence.
Dolichoderomorphs 100 105 120 Burmomyrma, a putative aneuretine in Burmese amber (Dlussky, 1996). Aneuretinae may
be paraphyletic relative to Dolichoderinae, so this fossil calibrates stem
dolichoderomorphs.
Formicinae 92 95 120 Kyromyrma in New Jersey amber (Grimaldi & Agosti, 2000).
Lasius 42 60 90 Lasius in Baltic amber (Dlussky, 2011).
Myrmeciinae 45 60 90 Archimyrmex (Myrmeciinae) in middle Eocene deposits of the United States, Argentine
and Germany (Dlussky, 2012).
Ponerinae 42 60 90 Platythyrea in Baltic amber (Wheeler, 1915).
Tatuidris 42 60 90 Agroecomyrmex duisburgi (Agroecomyrmecinae) in Baltic amber (Wheeler, 1915;
Dlussky & Rasnitsyn, 2009).

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


Phylogeny and evolution of myrmicine ants 67

Table 4. Adjacency matrix used in lagrange analysis, for the Most Pogonomyrmex species are noted for their granivory and
Neotropical (T), Nearctic (N), Palearctic (P), Afrotropical (E), Indo- preference for xeric environments (Cole, 1968; Taber, 1998;
malayan (O) and Australasian regions (A). Dispersal rates for adja- Johnson, 2001), but these are evidently derived traits within
cent areas are given for the time periods 0–50 Ma (above diagonal)
the Pogonomyrmecini. Species in the Pogonomyrmex angustus
and 50–100 Ma (below diagonal). Rates are estimated to be lower
complex and in the genus Hylomyrma are found mostly in mesic
(0.5) for adjacent regions separated by large water gaps, i.e. Neotropi-
cal/Afrotropical, Neotropical/Australian and Afrotropical/Australian in habitats and are not specialist seed-harvesters (Kempf, 1973;
the first time period, and Indomalayan/Australian in the second time Brown, 2000; P.S. Ward, personal observation).
period, based on broad-scale reconstructions of tectonic history over
these time periods (Scotese, 2010).
Clade 3: Stenammini
T N P E O A

T – 1.0 0.0 0.5 0.0 0.5 We recover, with strong support (PP 1.00 BS 100), a cluster of
N 1.0 – 1.0 0.0 0.0 0.0 taxa comprising the genera Stenamma, Messor, Aphaenogaster,
P 0.0 1.0 – 1.0 1.0 0.0 Goniomma and Oxyopomyrmex. This group, representing a
E 1.0 0.0 1.0 – 1.0 0.5 modified circumscription of the tribe Stenammini (Branstetter,
O 0.0 0.0 1.0 1.0 – 1.0 2012), is sister to all myrmicines except Myrmicini and Pogon-
A 1.0 0.0 0.0 1.0 0.5 – omyrmecini (PP 1.00 BS 76). Within Stenammini, Messor and
Aphaenogaster are demonstrably nonmonophyletic: ants in the
genus Messor consist of separate Old World and New World
Clade 1: Myrmicini clades (see also Brady et al., 2006), whereas those assigned
to the genus Aphaenogaster represent at least four different
The reciprocally monophyletic genera Manica and Myrmica
lineages. The localized Mediterranean taxa, Goniomma and
together form a clade (PP 1.00 BS 100) that is sister to all
Oxyopomyrmex, are one another’s closest relatives (see also
other Myrmicinae (PP 0.97 BS 78), a result that is consistent
Moreau et al., 2006); and the species-rich genus Stenamma is
with the generalized morphology of these ants (Dlussky &
recovered as sister to a New World clade composed of the erst-
Radchenko, 2009). We refer to this clade as the tribe Myrmicini,
while genera Novomessor (represented here by Aphaenogaster
restricting its membership to these two extant genera, and
albisetosa) and Veromessor (represented by Messor andrei). We
excluding the other five genera currently placed in the tribe. Two
resurrect Veromessor from synonymy (see Taxonomy, below),
of these, Pogonomyrmex and Hylomyrma, form a separate clade
but reclassification of the several Aphaenogaster-like lineages
(Clade 2), whereas the other three (Eutetramorium, Huberia and
awaits a more comprehensive investigation of the phylogeny of
Secostruma) belong to a large clade of predominantly Old World
the Stenammini, currently underway (M. Branstetter, personal
ants (Clade 6; see below). These results, and the relationships
communication).
among the four Myrmica species that we sampled, are in
agreement with other recent molecular phylogenetic analyses of
Myrmicini (Jansen & Savolainen, 2010; Jansen et al., 2010), but
Clade 4: Solenopsidini
our study is the first to establish its position as sister to all other
members of the subfamily.
This clade (PP 1.00 BS 99) is an amalgam of groups to which
In addition to their generalized morphology, Myrmica workers
we assign the tribal name Solenopsidini, one of two oldest
retain reproductive (haploid egg-laying) potential and are gen-
available names (the other is Myrmicariini). It includes the
eralist predators and scavengers (Seifert, 1988; Elmes, 1991;
genera Adelomyrmex, Anillomyrma, Austromorium, Baracidris,
Wardlaw & Elmes, 1998). Similar habits are thought to char-
Bariamyrma, Carebarella, Cryptomyrmex, Dolopomyrmex,
acterize the genus Manica, although less is known about the
Megalomyrmex, Monomorium, Myrmicaria, Oxyepoecus,
biology of these ants (Wheeler & Wheeler, 1970).
Rogeria, Solenopsis, Stegomyrmex, Tropidomyrmex and Tyran-
nomyrmex. The unsampled genus Bondroitia probably also
Clade 2: Pogonomyrmecini belongs here (Bolton, 1987). The genus Stegomyrmex is sister
to all other solenopsidines (PP 1.00 BS 100). According to our
The two genera Pogonomyrmex and Hylomyrma collectively findings, Rogeria, Adelomyrmex, Solenopsis and Monomorium
constitute a well-supported clade (PP 1.00 BS 100) that is sister are not monophyletic. In the case of Rogeria and Adelomyrmex
to all remaining myrmicines except Myrmicini (PP 1.00 BS it appears that the Asian-Pacific species of these genera repre-
98). We apply the name Pogonomyrmecini, trib.n. to this clade sent lineages separate from their Neotropical counterparts, but
(see Taxonomy section below). Hylomyrma is nested within additional taxon sampling is required to confirm this. Care-
Pogonomyrmex, but monophyly of the latter can be restored barella is nested phylogenetically within Solenopsis; hence
by creation of a new genus for Pogonomyrmex angustus and synonymization of the former under the latter (see Taxonomy,
its close relatives. These form the Pogonomyrmex angustus below) restores Solenopsis monophyly.
complex, a monophyletic group that is sister to all other Monomorium presents a much more complicated
pogonomyrmecines (C. Moreau and R.A. Johnson, personal situation – species assigned to this genus are scattered across
communication). the solenopsidine clade, and several are even positioned

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


68 P. S. Ward et al.

Strumigenys godeffroyi
Strumigenys chyzeri
Strumigenys emmae
Strumigenys chiricahua
0.96/69 Strumigenys ocypete
Strumigenys membranifera
Strumigenys exiguaevitae
Strumigenys olsoni
Strumigenys maxillaris
Strumigenys dicomas Carebara affinis
Strumigenys coveri
Strumigenys rogeri Carebara urichi
Strumigenys ludovici Carebara AU01
Strumigenys erynnes Carebara alperti
Strumigenys biolleyi Carebara nosindambo
Strumigenys gundlachi Carebara vidua
Strumigenys nitens Diplomorium longipenne
Strumigenys ambatrix Cardiocondyla thoracica
Pilotrochus besmerus Cardiocondyla MY01
Phalacromyrmex fugax 0.96/18 Cardiocondyla mauritanica
Talaridris mandibularis Ocymyrmex fortior cf
Eurhopalothrix gravis Cataulacus hispidulus
0.81/44 Cataulacus ebrardi
Eurhopalothrix australis
Octostruma EC01 Atopomyrmex mocquerysi
Basiceros manni Nesomyrmex MG08
Rhopalothrix isthmica Nesomyrmex MG01
Protalaridris armata Nesomyrmex madecassus
Pheidole longispinosa Nesomyrmex wilda
0.96/37 Machomyrma dispar Nesomyrmex echinatinodis
Anisopheidole antipodum Xenomyrmex floridanus
Pheidole pallidula Vitsika suspicax
Pheidole lucioccipitalis Myrmisaraka brevis
Pheidole hyatti Malagidris belti
Pheidole clydei Malagidris jugum
Pheidole rhea Royidris longiseta
Pheidole fimbriata Royidris notorthotenes
0.99/58 Cephalotes texanus Eutetramorium mocquerysi
Cephalotes atratus Metapone PG01
Procryptocerus scabriusculus Metapone madagascarica
0.98/16 Trachymyrmex arizonensis Liomyrmex gestroi
Atta texana Podomyrma silvicola
0.93/60 Acromyrmex versicolor Huberia brounii
Cyphomyrmex longiscapus cf Monomorium destructor
Cyphomyrmex cornutus Monomorium criniceps
Mycetosoritis hartmanni Monomorium robustior
Mycetarotes acutus Dacetinops ignotus cf
Kalathomyrmex emeryi Crematogaster emeryana
Myrmicocrypta infuscata cf Crematogaster kelleri
Mycocepurus goeldii Crematogaster modiglianii
Apterostigma auriculatum Crematogaster sordidula
0.93/36 0.93/18 Crematogaster acuta
Mesostruma browni
1.00/59 Epopostruma monstrosa Meranoplus radamae
Colobostruma unicorna Meranoplus castaneus
1.00/60 Orectognathus versicolor Pristomyrmex punctatus
0.95/49 Microdaceton tibialis 1.00/65 Pristomyrmex orbiceps
Lenomyrmex colwelli Perissomyrmex snyderi
Daceton armigerum Secostruma MY01
0.96/41 Dilobocondyla borneensis cf
Acanthognathus ocellatus
Wasmannia auropunctata 0.91/48 Recurvidris TH01
Allomerus octoarticulatus Adlerzia froggatti
Blepharidatta conops Lophomyrmex ambiguus
Lachnomyrmex scrobiculatus Lordomyrma desupra
Diaphoromyrma sofiae Lordomyrma bhutanensis cf
Tranopelta gilva Propodilobus pingorum
Tranopelta subterranea Ancyridris polyrhachioides nr
Ochetomyrmex semipolitus Mayriella ebbei
Ochetomyrmex neopolitus Acanthomyrmex ferox
Attini Monomorium MG04
Monomorium exiguum
Myrmecina graminicola
Myrmecina americana
Monomorium fieldi nr Myrmecina PG01
Monomorium ergatogyna Rotastruma stenoceps
Monomorium madecassum Rotastruma recava
Monomorium pharaonis Romblonella scrobifera
Monomorium hanneli Poecilomyrma myrmecodiae
Cryptomyrmex boltoni Paratopula TH01
Adelomyrmex SC01 Proatta butteli
Baracidris sitra Dacatria templaris
Adelomyrmex paratristani Tetramorium spinosum
Monomorium fisheri Tetramorium severini
Monomorium ferodens Tetramorium MG125
Monomorium sechellense Rhoptromyrmex transversinodis
Myrmicaria exigua Teleutomyrmex kutteri
Myrmicaria carinata Anergates atratulus
0.94/40 0.90/20 Strongylognathus testaceus
Anillomyrma decamera
Anillomyrma AFRC TZ 01 Tetramorium caespitum cf
Monomorium latinode Tetramorium validiusculum
Monomorium brocha Tetramorium venator
Tyrannomyrmex legatus Tetramorium taylori
Megalomyrmex silvestrii Tetramorium CF01
Oxyepoecus vezenyii 1.00/48
Vollenhovia emeryi
Monomorium denticulatum Myrmicine genus26 PH02
Monomorium antarcticum 0.97/23 Calyptomyrmex kaurus
Austromorium flavigaster Dicroaspis KM01
Solenopsis succinea Cyphoidris exalta
Solenopsis molesta 0.99/17 Kartidris sparsipila
Solenopsis xyloni Terataner MG27
Solenopsis mameti Terataner MG02
Solenopsis ZA02 1.00/54 Terataner bottegoi
0.99/43 Solenopsis fugax Rhopalomastix rothneyi
Solenopsis papuana Melissotarsus insularis
Carebarella PA01 Myrmoxenus bernardi
Tropidomyrmex elianae Chalepoxenus kutteri
Kempfidris inusualis Temnothorax rugatulus
Rogeria stigmatica Temnothorax poeyi
1.00/67 Leptothorax muscorum complex
Bariamyrma hispidula
Rogeria nevadensis Formicoxenus diversipilosus
0.99/48 Harpagoxenus sublaevis
Dolopomyrmex pilatus
Stegomyrmex manni Vombisidris bilongrudi
Solenopsidini Messor wasmanni
Messor denticornis
Gauromyrmex acanthinus
Tetheamyrma subspongia
Aphaenogaster occidentalis
Oxyopomyrmex santschii
Crematogastrini
Goniomma blanci
Aphaenogaster araneoides
Aphaenogaster swammerdami
Aphaenogaster pythia
Stenamma striatulum
Stenammini Stenamma dyscheres
Stenamma expolitum
Messor andrei
Aphaenogaster albisetosa
Pogonomyrmex vermiculatus
Pogonomyrmex subdentatus
Pogonomyrmex imberbiculus
Pogonomyrmex striatinodus
Hylomyrma blandiens
PogonomyrmeciniMyrmica tahoensis Pogonomyrmex angustus
Myrmica striolagaster
Myrmica rugiventris
Myrmica kotokui
Manica rubida
Myrmicini Manica bradleyi 0.01 length units

0.91/64

Formicoid Rhytidoponera chalybaea


Acanthoponera minor
Myrmelachista flavocotea
Lasius californicus
Nothomyrmecia macrops
Myrmecia pyriformis
Tetraponera rufonigra
Dolichoderus pustulatus
Aneuretus simoni
Tatuidris tatusia_2
Poneroid Tatuidris tatusia
Ankylomyrma coronacantha
Platythyrea mocquerysi
Paraponera clavata
Proceratium stictum
1.00/62 Stigmatomma pallipes
Amblyopone australis
Apterogyna ZA01

Fig. 1. Maximum likelihood (ML) phylogeny of the ant subfamily Myrmicinae, including formicoid and poneroid outgroups, estimated with a
partitioned ML analysis of a concatenated 11-gene dataset (−lnL = 226 779.1302). Support values were generated in a separate partitioned ML bootstrap
analysis with 1000 replicates, and a separate partitioned Bayesian analysis. Node support is indicated by the following colour scheme (PP, Bayesian
posterior probability; BS, ML bootstrap percentage): red dots PP = 1.00, BS >90; brown dots PP = 1.00, BS = 70–90; blue dots PP = 0.95–0.99, BS
≥70. For nodes of interest that do not fall into these three categories, we indicate actual support values as PP/BS.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


Phylogeny and evolution of myrmicine ants 69

Table 5. Support values under different modes of analysis, and crown age estimates from the beast analysis, for selected clades of Myrmicinae.

Cladea BS PP PP (i) PP (ii) PP (iii) PP (iv) PP (beast) Age (Ma) 95% HPD

Myrmicinae 98 1.00 1.00 1.00 1.00 1.00 1.00 98.6 87.9–109.6


Myrmicini 100 1.00 1.00 1.00 1.00 1.00 1.00 52.3 45.3–61.0
All myrmicines except Myrmicini 78 0.97 1.00 0.99 0.95 0.94 0.92 95.4 85.2–106.0
Pogonomyrmecini 100 1.00 1.00 1.00 1.00 1.00 1.00 54.3 43.7–66.1
Stenammini 100 1.00 1.00 1.00 1.00 1.00 1.00 52.9 48.1–58.6
Solenopsidini 99 1.00 1.00 1.00 1.00 1.00 1.00 71.1 62.4–79.7
Attini 100 1.00 1.00 1.00 1.00 1.00 1.00 67.0 59.2–75.4
Crematogastrini 98 1.00 1.00 1.00 1.00 1.00 1.00 70.8 64.1–77.9
Strumigenys + Phalacromyrmex genus-group 100 1.00 1.00 1.00 1.00 1.00 1.00 48.4 40.4–56.5
Strumigenys + Phalacro. genus-group + Basiceros genus-group 98 1.00 1.00 1.00 1.00 1.00 1.00 55.9 48.3–63.6
Daceton genus-group 41 0.96 1.00 1.00 0.87 – 1.00 58.9 51.3–67.0
Atta genus-group 95 1.00 1.00 1.00 1.00 1.00 1.00 55.6 47.9–63.5
Atta genus-group + Daceton genus-group 59 1.00 0.69 0.63 0.86 – 1.00 59.9 52.3–67.8
Pheidole + Cephalotes genus-group 58 0.99 – – 0.96 0.99 1.00 58.0 49.8–66.4
Cardiocondyla + Ocymyrmex 18 0.96 0.76 0.62 0.93 – 0.74 50.7 43.3–58.0
Core formicoxenines (including Gauromyrmex) 93 1.00 1.00 1.00 1.00 1.00 1.00 52.7 46.7–59.3
Tetramorium s.l. 100 1.00 1.00 1.00 1.00 1.00 1.00 20.1 15.4–24.8
Eutetramorium genus-group 92 1.00 – – 1.00 1.00 1.00 46.4 37.8–55.5
‘Monomorium’ scabriceps- and destructor-groups 100 1.00 1.00 1.00 1.00 1.00 1.00 16.2 9.3–23.8
Lordomyrma + Ancyridris + Propodilobus 100 1.00 1.00 1.00 1.00 1.00 1.00 28.9 18.9–39.8
Paratopula + Poecilomyrma + Romblonella + Rotastruma 70 1.00 1.00 1.00 1.00 1.00 1.00 50.8 39.2–62.0
a Tribalnames are used in the new sense adopted in this paper. The Phalacromyrmex, Basiceros, Daceton, Atta and Cephalotes genus-groups refer to
the erstwhile tribes Phalacromyrmecini, Basicerotini, Dacetini, Attini and Cephalotini, respectively.
BS = maximum likelihood bootstrap proportion from concatenated partitioned analysis of the full dataset; PP = Bayesian posterior probability from
concatenated, partitioned Bayesian analysis of the full dataset; PP (i) to PP (iv) = Bayesian posterior probabilities from treatments (i) to (iv), respectively,
addressing base frequency heterogeneity (see text); PP (beast) = Bayesian posterior probability from the beast analysis. PP values less than 0.50 are
not recorded. Age is estimated crown-group age in millions of years; 95% HPD is 95% highest probability density.

outside it (see discussion of Clade 6 below). There are two Clade 5: Attini (new sense)
pairs of austral clades, for example, comprising Oxyepoe-
cus + ‘Monomorium’ denticulatum (PP 1.00 BS 100) and This well-supported clade (PP 1.00 BS 100) comprises
Austromorium + ‘Monomorium’ antarcticum (PP 1.00 BS 100). a diverse selection of predominantly Neotropical taxa, for
In the Paleotropics ‘Monomorium’ brocha and ‘Monomorium’ which the oldest available tribal name is Attini. This clade
latinode are sister to Tyrannomyrmex (PP 1.00 BS 94). There contains Allomerus, Anisopheidole, Blepharidatta, Cephalotes,
is, however, a core clade of Monomorium s.s., corresponding Diaphoromyrma, Lachnomyrmex, Lenomyrmex, Machomyrma,
to the monomorium- and salomonis-groups of Bolton (1987), Ochetomyrmex, Pheidole, Procryptocerus, Tranopelta and Was-
mannia, as well as all the genera currently assigned to the
that is strongly supported as a monophyletic group (PP 1.00 BS
tribes Attini (i.e. the fungus-growing ants), Basicerotini, Dace-
100). We also recover, with moderate support (PP 1.00 BS 72),
tini and Phalacromyrmecini. In the interests of establishing a
a more inclusive clade that encompasses the hanneli-, hilde-
stable, phylogenetic classification of the Myrmicinae, devoid
brandti-, monomorium- and salomonis-groups of Monomorium
of paraphyletic groups, we subsume these taxa under a single
(Bolton, 1987; Heterick, 2006), but also includes the genera
tribe, encompassing all members of Clade 5 (see Taxonomy
Adelomyrmex, Anillomyrma, Baracidris, Cryptomyrmex and
section below). Although Attini is the oldest available name we
Myrmicaria. We reinstate the genus Syllophopsis for the mem- recognize that within a large body of biological literature the
bers of the Monomorium hildebrandti-group (see Taxonomy noun ‘Attini’ and the adjective ‘attine’ refer exclusively to the
section below). Monophyly of the hildebrandti-group is only fungus-growing ants. To avoid upsetting general usage by per-
weakly supported in our analyses (PP 66, BS 43) but the group manently expanding the definition of Attini, we are in the pro-
has distinctive morphological features (Heterick, 2006; B. cess of proposing to the International Commission on Zoological
Bolton, personal communication) that indicate monophyly. Nomenclature the substitution of the junior tribal synonym Phei-
Several genera currently placed in Solenopsidini do not belong dolini for Attini.
in this clade: Allomerus, Machomyrma and Tranopelta are We uncover several intriguing relationships within this clade.
in Clade 5, whereas Adlerzia, Carebara, Diplomorium and The trap-jaw ants of the old tribe Dacetini are not monophyletic:
Mayriella are in Clade 6 (Fig. 1). Based on the original species placed in Strumigenys form a separate clade that is
description (Terayama, 2009) Formosimyrma, which we did not sister to phalacromyrmecine ants, represented in our study
sequence, is probably also a member of Clade 6. by Phalacromyrmex and Pilotrochus. The phalacromyrmecines

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


70 P. S. Ward et al.

and Strumigenys are in turn sister to basicerotine ants. This analysis of the full dataset these two are sister taxa (PP 0.96 BS
arrangement receives strong support (PP 1.00 BS 98–100) at the 18). However, in treatment (iv), the Bayesian analysis address-
relevant nodes. Species of long-mandibulate Strumigenys with ing base frequency heterogeneity in which the 10% of characters
kinetic action are nested within groups of short-mandibulate deviating most strongly from observed average base frequencies
species formerly assigned to the genus Pyramica. This justifies were removed, the two genera are well separated: Cardiocondyla
the synonymy of the latter under the former (Baroni Urbani & is in a weakly supported clade (PP 0.81) containing Tetramor-
De Andrade, 2007), and supports the hypothesis (Bolton, 1999) ium and several other genera, whereas Ocymyrmex appears in a
that kinetic mandibles are a derived state in the genus. moderately well supported clade (PP 0.94) with Meranoplus and
The true dacetine ants (PP 96 BS 41) comprise all genera Nesomyrmex. We focus in this discussion on other subgroups
currently placed in this group except Strumigenys, but they also within Clade 6 that we recover more consistently and with more
include the enigmatic genus Lenomyrmex, whose affinities were robust support.
previously unclear (Fernández & Palacio, 1999). Dacetine ants One of these is a clade of core formicoxenines (PP 1.00
are sister to the fungus-growing ants, hereafter referred to as the BS 93) comprising Gauromyrmex, Vombisidris, Harpagoxenus,
Atta genus-group (PP 1.00 BS 59 for the larger clade; PP 1.00 Formicoxenus, Leptothorax, Temnothorax, Chalepoxenus and
BS 95 for the Atta genus-group). Previous analyses (Schultz Myrmoxenus, with the last two nested within Temnothorax.
& Brady, 2008) also hinted at a relationship between the Atta Based on mtDNA sequence data (Beibl et al., 2005) Protomo-
genus-group and dacetine ants, but without significant support. gnathus also falls within Temnothorax. Most of the other genera
Our results motivate a search for features in common between currently placed in Formicoxenini, including Atopomyrmex,
these two groups that might provide insight into the origins of Dilobocondyla, Ochetomyrmex, Nesomyrmex, Podomyrma and
specialized biology in both clades. Terataner, show no close relationship to this group (Fig. 1).
Elsewhere in the Attini (s.l.) we find that the large genus Because we cannot reliably place Cardiocondyla, the question
Pheidole is nonmonophyletic because two derivative genera, of its possible relationship to the core formicoxenines remains
Machomyrma and Anisopheidole, are nested within it. Syn- unresolved.
onymy of these two under Pheidole (see Taxonomy section The tetramoriine ants are a very strongly supported group (PP
below) restores monophyly (PP 1.00 BS 100). The genus Chi- 1.00, BS 100), comprising Tetramorium and derivative ‘satellite
maeridris (Wilson, 1989) is probably also nested within Phei- genera’ that are nested within it: Rhoptromyrmex, Anergates,
dole, but we were unable to sequence this ant, so its status Teleutomyrmex and Strongylognathus. These are social parasites
remains uncertain. The species of Pheidole that we sampled whose divergent morphology caused them to be placed in their
from the Old World, including Pheidole pallidula, form a clade own genera. We identify a clade (PP 1.00 BS 100) composed of
(PP 1.00 BS 65) nested within a paraphyletic group of New Calyptomyrmex, Vollenhovia and an undescribed genus from the
World Pheidole, supporting the hypothesis of a New World ori- Philippines, as sister to the tetramoriines, with moderate support
gin of the genus and a single dispersal event to the Old World (cf. (PP 1.00 BS 86).
Moreau, 2008), although the recent report of Pheidole in Baltic Three groups of species assigned, or recently assigned,
amber (Dubovikoff, 2011) suggests the possibility of another to Monomorium also appear in Crematogastrini. The genus
earlier colonization, possibly by a stem Pheidole. The sister Royidris (PP 1.00 BS 100), formerly the ‘Monomorium’
group of Pheidole appears to be the turtle ants, or Cephalotes shuckardi-group (Heterick, 2006), is part of a well-supported
genus-group (PP 0.99 BS 58 for the more inclusive clade, PP clade (PP 1.00 BS 92) of morphologically disparate genera
1.00 BS 100 for the Cephalotes genus-group), a rather surpris- all endemic to Madagascar, the Eutetramorium genus-group
ing finding in view of the morphological discrepancy between (Bolton & Fisher, 2014). The ‘Monomorium’ scabriceps- and
these two groups. destructor-groups form their own clade (PP 1.00 BS 100) that
is part of a larger group (PP 1.00 BS 92) comprising two other
clades: Huberia + Podomyrma + Liomyrmex + Metapone (PP
Clade 6: Crematogastrini (new sense) 1.00 BS 99) and the aforementioned Eutetramorium cluster. We
resurrect the genus Trichomyrmex to contain the scabriceps-
The sixth clade (PP 1.00 BS 98) is also a highly diverse assem- and destructor-groups (see below).
blage, centred primarily in the Old World tropics. We apply one Lordomyrma + Ancyridris + Propodilobus form a very
of the two oldest tribal names, Crematogastrini, to this group strongly supported clade (PP 1.00 BS 100), with very short
(the other available name is Formicoxenini). It contains all the branches separating these three genera. The sister group of
remaining myrmicine genera, including Cardiocondyla, Care- this diverse Australasian clade (Lucky & Sarnat, 2010) is not
bara, Cataulacus, Crematogaster, Leptothorax, Lordomyrma, confidently identified in our analyses, although we can reject
Meranoplus, Myrmecina, Nesomyrmex, Podomyrma, Temnotho- some candidate taxa, namely Bariamyrma, Lachnomyrmex
rax, Tetramorium, Vollenhovia and about 50 other genera. and Rogeria (cf. Taylor, 2009), because these are in distant
Relationships are not well resolved at the base of this clade, clades. We do not find evidence of a close relationship of the
suggesting a period of rapid diversification early in its history. Lordomyrma clade to the genera Cyphoidris or Dacetinops
An additional challenge is posed by the long-branched taxa within the Crematogastrini, but attention should be focused
Ocymyrmex and Cardiocondyla (Fig. 1), whose positions are on Indomyrma (Brown, 1986) and Lasiomyrma (Terayama &
unstable across different analytical treatments. In the Bayesian Yamane, 2000), two taxa that we were unable to sequence.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


Phylogeny and evolution of myrmicine ants 71

The Indo-Pacific genera Paratopula, Poecilomyrma, attributed to this subfamily by Dlussky et al. (2004), but this
Romblonella and Rotastruma constitute a clade (PP 1.00 has been questioned by others (Wilson & Hölldobler, 2005;
BS 70) in which Paratopula is sister to the other three Archibald et al., 2006) and thus Afromyrma was not used as an
(PP 1.00 BS 99). Among other findings, we also recover a priori calibration in our dating analysis. Otherwise the first
the following sister-group relationships in Crematogas- myrmicine ants are not reported until early Eocene (50–55 Ma)
trini: Melissotarsus + Rhopalomastix (PP 1.00 BS 100), amber deposits from China (Hong, 2002), India (Rust et al.,
Cyphoidris + Dicroaspis (PP 1.00 BS 75), Dacatria + Proatta 2010) and France (Aria et al., 2011). These await detailed
(PP 1.00 BS 75), Dilobocondyla + Secostruma (PP 1.00 BS study – or re-evaluation in the case of the Chinese amber
100) and Diplomorium + Carebara (PP 1.00 BS 99). Carebara inclusions (LaPolla et al., 2013) – but they reveal considerable
affinis, a representative of the erstwhile genus Pheidologeton, taxonomic diversity, consistent with our inference that the major
is nested well within Carebara (Fig. 1), justifying synonymy of lineages of Myrmicinae had already originated by this time.
the former under the latter (Fischer et al., 2014). Our divergence date estimates indicate that the early and
The affinities of the large and diverse genus Crematogaster are mid-Eocene (55–40 Ma) was an important period of diversifi-
uncertain (Blaimer, 2012b). We obtain weak evidence (PP 93 BS cation for myrmicine ants (Table 5). Among taxa with estimated
18) that its sister group is the Paleotropical genus Meranoplus crown origins at about this time are the three aforementioned
and that Recurvidris, previously hypothesized to be the closest tribes (Myrmicini, Pogonomyrmecini, Stenammini), and numer-
relative of Crematogaster (Bolton, 2003), is sister to Adlerzia ous other suprageneric taxa including the following (ages in Ma,
(PP 91 BS 48). with 95% HPD): Daceton genus-group (58.9; 51.3–67.0), Atta
There is also modest support for a Peris- genus-group (55.6; 47.9–63.5), Cephalotes genus-group (43.0;
somyrmex + Pristomyrmex sister pair (PP 1.00 BS 65), but 33.6–52.6), Basiceros genus-group (45.0; 36.8–53.4) and ‘core
surprisingly we do not recover a more inclusive clade con- formicoxenines’, as defined above (52.7; 46.7–59.3). Our age
taining these two plus Myrmecina and Acanthomyrmex: the inferences for the fungus-growing ants (Atta genus-group) are
erstwhile tribe Myrmecinini (Ogata & Okido, 2007). Nev- comparable to those of a previous Bayesian dating analysis of
ertheless, we cannot confidently reject this putative clade, the group (Schultz & Brady, 2008; 52 Ma, 95% HPD 44–59).
because Myrmecina and Acanthomyrmex are isolated taxa of Nevertheless we find that some of the most prominent con-
uncertain position in our tree, emerging from the bushy base of temporary genera in the Myrmicinae have relatively recent
Crematogastrini. Other apparently isolated genera, not strongly crown-group origins. Examples include (ages in Ma, with
associated with particular taxa, include Dacetinops, Kartidris, 95% HPD) Solenopsis (39.1; 31.4–47.1), Pheidole (35.2;
Lophomyrmex, Mayriella, Nesomyrmex, Tetheamyrma and 24.9–46.6), Cephalotes (25.3; 19.5–31.9), Strumigenys (33.1;
Xenomyrmex. There is a weak indication that Tetheamyrma is 27.2–39.2), Crematogaster (34.7; 28.1–42.4) and Tetramorium
sister to all other Crematogastrini (PP 90 BS 20), but confir- (20.1; 15.4–24.8). These are highly successful, species-rich
mation of this and other relationships within Crematogastrini genera that appear to have undergone intense diversification
requires improved taxon sampling and, more critically, a much since the Eocene. Our estimates of the crown ages of Phei-
larger sampling of genes. dole and Cephalotes are considerably younger than those of
other recent studies (Pheidole: 58.4–61.2 Ma, Moreau, 2008;
Cephalotes: 46 Ma, 95% HPD 37–56 Ma, Price et al., 2014),
Divergence dating and biogeographic history suggesting a more compressed history of diversification in these
groups than inferred by those investigators.
The chronogram derived from a beast analysis (Fig. 2) recovers Results of the lagrange analysis show that the two largest
a crown-group age for the Myrmicinae of 98.6 Ma [95% high- clades of Myrmicinae, the sister taxa Attini and Crematogas-
est posterior density (HPD) 87.9–109.6 Ma]. The major clades trini, originated and diversified largely in the Neotropics and
within the subfamily are considerably younger, with estimated Paleotropics, respectively (Fig. 3, Table S7). Some Crematogas-
crown ages of 52.3 Ma for Myrmicini, 54.3 Ma for Pogono- trini lineages subsequently spread into the Palearctic region and
myrmecini, 52.9 Ma for Stenammini, 71.1 Ma for Solenopsidini, Australia, and a few (such as the genera Nesomyrmex, Temnotho-
67.0 Ma for Attini and 70.8 Ma for Crematogastrini (Table 5). It rax, Carebara and Crematogaster) colonized the New World.
is striking that the ‘basal’ (i.e. earliest branching) lineages of Conversely, the early evolution of Attini was centred in the
myrmicines (Myrmicini, Pogonomyrmecini, Stenammini) have New World tropics, with subsequent dispersal events, in the late
younger crown ages than the other major clades, attesting to sub- Eocene, to the Afrotropics (Strumigenys) and Australia (Dace-
stantial pruning of the stem lineages of these three groups. ton genus-group, Eurhopalothrix). Dispersal to the Old World
Our inferred crown age of ∼100 Ma for Myrmicinae is by Pheidole is estimated to have occurred more recently, in the
somewhat older than estimates from other studies, which are middle Miocene (Fig. 3).
in the range of 75–90 Ma (Brady et al., 2006; Moreau & The Solenopsidini is another clade which is inferred to have
Bell, 2013; Schmidt, 2013), although the 95% HPD values had a Neotropical origin (Table S7). In contrast to the Attini,
overlap with those of most estimates in Moreau & Bell (2013). however, there appears to have been an early colonization
The fossil record does not include any definitive Myrmicinae of the Paleotropics about 50 Ma, which led to substantial
from the Cretaceous. Afromyrma petrosa, described from a diversification there. There was also an independent dispersal
Turonian mudstone imprint from Orapa, Botswana (91 Ma), was of solenopsidines from the Neotropics to Australia, presumably

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


72 P. S. Ward et al.

Crematogastrini

Strumigenys chyzeri
Strumigenys godeffroyi
Strumigenys emmae
Strumigenys chiricahua
Strumigenys ocypete
Strumigenys membranifera
Strumigenys exiguaevitae
Strumigenys olsoni
Strumigenys maxillaris

Attini
Strumigenys coveri
Strumigenys dicomas
Strumigenys rogeri
Strumigenys erynnes
Strumigenys ludovici
Strumigenys gundlachi
Strumigenys biolleyi
Strumigenys nitens
Strumigenys ambatrix
Pilotrochus besmerus
Phalacromyrmex fugax
Eurhopalothrix gravis
Talaridris mandibularis
Eurhopalothrix australis
Octostruma EC01
Basiceros manni
Rhopalothrix isthmica
Protalaridris armata
Machomyrma dispar
Pheidole longispinosa
Anisopheidole antipodum
Pheidole lucioccipitalis
Pheidole pallidula
Pheidole hyatti
Pheidole clydei
Pheidole rhea
Pheidole fimbriata
Cephalotes atratus
Cephalotes texanus
Procryptocerus scabriusculus
Trachymyrmex arizonensis
Acromyrmex versicolor
Atta texana
Cyphomyrmex longiscapus cf
Cyphomyrmex cornutus
Mycetosoritis hartmanni
Mycetarotes acutus
Kalathomyrmex emeryi
Mycocepurus goeldii
Myrmicocrypta infuscata cf
Apterostigma auriculatum
Epopostruma monstrosa
Mesostruma browni
Colobostruma unicorna
Orectognathus versicolor
Microdaceton tibialis
Daceton armigerum
Lenomyrmex colwelli
Acanthognathus ocellatus
Allomerus octoarticulatus
Wasmannia auropunctata
Blepharidatta conops
Lachnomyrmex scrobiculatus
Diaphoromyrma sofiae
Tranopelta subterranea
Tranopelta gilva
Ochetomyrmex neopolitus
Ochetomyrmex semipolitus
Monomorium ergatogyna
Monomorium fieldi nr
Monomorium exiguum
Monomorium MG04
Monomorium madecassum
Monomorium pharaonis
Monomorium hanneli
Solenopsidini

Adelomyrmex SC01
Cryptomyrmex boltoni
Baracidris sitra
Adelomyrmex paratristani
Monomorium ferodens
Monomorium fisheri
Monomorium sechellense
Anillomyrma decamera
Anillomyrma AFRC TZ 01
Myrmicaria carinata
Myrmicaria exigua
Monomorium brocha
Monomorium latinode
Tyrannomyrmex legatus
Solenopsis molesta
Solenopsis succinea
Solenopsis xyloni
Solenopsis mameti
Solenopsis fugax
Solenopsis ZA02
Solenopsis papuana
Carebarella PA01
Kempfidris inusualis
Tropidomyrmex elianae
Austromorium flavigaster
Monomorium antarcticum
Monomorium denticulatum
Oxyepoecus vezenyii
Megalomyrmex silvestrii
Rogeria stigmatica
Bariamyrma hispidula
Dolopomyrmex pilatus
Rogeria nevadensis
Stegomyrmex manni
Messor denticornis
Messor wasmanni
Aphaenogaster occidentalis
Goniomma blanci
Oxyopomyrmex santschii

Stenammini Aphaenogaster araneoides


Aphaenogaster pythia
Aphaenogaster swammerdami
Stenamma striatulum
Stenamma dyscheres
Stenamma expolitum
Aphaenogaster albisetosa
Messor andrei
Pogonomyrmex subdentatus
Pogonomyrmex vermiculatus
Pogonomyrmecini
m Pogonomyrmex imberbiculus
Pogonomyrmex striatinodus
Hylomyrma blandiens
Pogonomyrmex angustus
Myrmica striolagaster
Myrmicini Myrmica tahoensis
Myrmica rugiventris
Myrmica kotokui
Manica bradleyi
Manica rubida

100 75 50 25 0
million years ago

Fig. 2. (a, b) Chronogram of Myrmicinae, from a beast analysis, with outgroups not shown. Blue bars depict the 95% HPD (highest probability
density), and are given for all nodes for which the 95% HPD function encompasses an estimate of 25 Ma or older.

via Antarctica, about 40–50 Ma (Fig. 3). The latter event was Of the three earliest diverging (‘basal’) lineages of Myr-
followed by a large radiation of Monomorium-like species micinae, the Stenammini are most abundant in the northern
in Australia (Ettershank, 1966; Heterick, 2001). The genus hemisphere and are most likely of Nearctic origin, although
Solenopsis itself is of Neotropical origin but has come to occupy selected lineages have dispersed to the northern Neotrop-
all major biogeographic regions; in this case dispersal outside ics and the Paleotropics; the Pogonomyrmecini arose in the
the Neotropics occurred more recently, within the last 25–30 Ma Neotropics and subsequently colonized the Nearctic region;
(Fig. 3). and the Myrmicini are reconstructed as having a Nearctic

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


Phylogeny and evolution of myrmicine ants 73

Cardiocondyla MY01
Cardiocondyla thoracica
Cardiocondyla mauritanica
Ocymyrmex fortior cf
Cataulacus ebrardi
Cataulacus hispidulus
Atopomyrmex mocquerysi
Nesomyrmex MG08
Nesomyrmex MG01
Nesomyrmex madecassus
Nesomyrmex echinatinodis
Nesomyrmex wilda
Xenomyrmex floridanus
Carebara alperti
Carebara AU01
Carebara affinis
Carebara urichi
Carebara nosindambo
Carebara vidua
Diplomorium longipenne
Vitsika suspicax
Myrmisaraka brevis
Malagidris jugum
Malagidris belti
Royidris notorthotenes
Royidris longiseta
Eutetramorium mocquerysi
Metapone madagascarica
Metapone PG01
Liomyrmex gestroi
Podomyrma silvicola
Huberia brounii
Monomorium criniceps
Monomorium destructor
Monomorium robustior
Dacetinops ignotus cf
Crematogaster emeryana
Crematogaster kelleri
Crematogaster modiglianii
Crematogaster acuta
Crematogaster sordidula
Meranoplus castaneus
Meranoplus radamae
Dilobocondyla borneensis cf
Secostruma MY01
Crematogastrini Pristomyrmex orbiceps
Pristomyrmex punctatus
Perissomyrmex snyderi
Recurvidris TH01
Adlerzia froggatti
Lophomyrmex ambiguus
Lordomyrma bhutanensis cf
Lordomyrma desupra
Ancyridris polyrhachioides nr
Propodilobus pingorum
Acanthomyrmex ferox
Mayriella ebbei
Myrmecina americana
Myrmecina graminicola
Myrmecina PG01
Dacatria templaris
Proatta butteli
Tetramorium severini
Tetramorium spinosum
Tetramorium MG125
Rhoptromyrmex transversinodis
Anergates atratulus
Teleutomyrmex kutteri
Strongylognathus testaceus
Tetramorium caespitum cf
Tetramorium venator
Tetramorium validiusculum
Tetramorium CF01
Tetramorium taylori
Myrmicine genus26 PH02
Vollenhovia emeryi
Calyptomyrmex kaurus
Cyphoidris exalta
Dicroaspis KM01
Kartidris sparsipila
Terataner MG02
Terataner MG27
Terataner bottegoi
Melissotarsus insularis
Rhopalomastix rothneyi
Chalepoxenus kutteri
Myrmoxenus bernardi
Temnothorax rugatulus
Temnothorax poeyi

other Myrmicinae Leptothorax muscorum complex


Formicoxenus diversipilosus
Harpagoxenus sublaevis
Vombisidris bilongrudi
Gauromyrmex acanthinus
Rotastruma recava
Rotastruma stenoceps
Romblonella scrobifera
Poecilomyrma myrmecodiae
Paratopula TH01
Tetheamyrma subspongia

100 75 50 25 0
million years ago

Fig. 2. Continued.

origin, with multiple dispersals to the Palearctic region (Fig. 3, combinations implicitly include the junior synonyms of the
Table S7). species names listed below. Author and year of publication
The two descendant branches of crown Myrmicinae, leading for all genus and species names can be found in AntCat
to (i) Myrmicini and (ii) all other myrmicines, are reconstructed (http://antcat.org/).
as most likely inheriting a Nearctic and Neotropical distribution, Veromessor stat.r. is resurrected from synonymy under
respectively (Table S7). The evolution of the ‘all other myr- Messor, generating the following revived combinations in
micines’ clade in the Late Cretaceous and early Tertiary appears, Veromessor: andrei, chamberlini, chicoensis, julianus, lariversi,
on the one hand, to have been concentrated in the Neotropics lobognathus, pergandei, smithi and stoddardi.
(Fig. 3), attesting to the importance of this region for early ant Syllophopsis stat.r. is resurrected from synonymy under
evolution (Moreau & Bell, 2013). On the other hand, dispersal Monomorium, to contain the Monomorium species in the
to other regions and diversification therein – starting as early as hildebrandti-group (sensu Heterick, 2006; this includes the
70 Ma for the Crematogastrini – has also been an integral part of fossulatum-group of Bolton, 1987). Ireneidris is removed
the myrmicine success story.
from synonymy under Monomorium, and newly synonymized
(syn.n.) under Syllophopsis. This creates the following new
combinations in Syllophopsis: adiastolon, aureorugosa, aus-
Taxonomy
tralica, dentata, ferodens, fisheri, gongromos, hildebrandti,
Generic changes infusca, kondratieffi, malamixta, sechellensis, sersalata, sub-
coeca, thrascolepta and vitiensis; and the following revived
The following generic changes are instituted, primarily to combinations in Syllophopsis: arnoldi (= jonesi, unnecessary
eliminate obvious cases of nonmonophyly. New and revived replacement name), cryptobia, elgonensis and modesta.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


74 P. S. Ward et al.

Crematogastrini

Nearctic
Neotropical Strumigenys chyzeri
Strumigenys godeffroyi
Strumigenys emmae

Australasian Strumigenys chiricahua


Strumigenys ocypete
Strumigenys membranifera
Strumigenys exiguaevitae
Indomalayan

Attini
Strumigenys olsoni
Strumigenys maxillaris
Strumigenys coveri

Afrotropical Strumigenys dicomas


Strumigenys rogeri
Strumigenys erynnes
Strumigenys ludovici
Palearctic Strumigenys gundlachi
Strumigenys biolleyi
Strumigenys nitens
Strumigenys ambatrix
Pilotrochus besmerus
Phalacromyrmex fugax
Eurhopalothrix gravis
Talaridris mandibularis
Eurhopalothrix australis
Octostruma EC01
Basiceros manni
Rhopalothrix isthmica
Protalaridris armata
Machomyrma dispar
Pheidole longispinosa
Anisopheidole antipodum
Pheidole lucioccipitalis
Pheidole pallidula
Pheidole hyatti
Pheidole clydei
Pheidole rhea
Pheidole fimbriata
Cephalotes atratus
Cephalotes texanus
Procryptocerus scabriusculus
Trachymyrmex arizonensis
Acromyrmex versicolor
Atta texana
Cyphomyrmex longiscapus cf
Cyphomyrmex cornutus
Mycetosoritis hartmanni
Mycetarotes acutus
Kalathomyrmex emeryi
Mycocepurus goeldii
Myrmicocrypta infuscata cf
Apterostigma auriculatum
Epopostruma monstrosa
Mesostruma browni
Colobostruma unicorna
Orectognathus versicolor
Microdaceton tibialis
Daceton armigerum
Lenomyrmex colwelli
Acanthognathus ocellatus
Allomerus octoarticulatus
Wasmannia auropunctata
Blepharidatta conops
Lachnomyrmex scrobiculatus
Diaphoromyrma sofiae
Tranopelta subterranea
Tranopelta gilva
Ochetomyrmex neopolitus
Ochetomyrmex semipolitus
Monomorium ergatogyna
Monomorium fieldi nr
Monomorium exiguum
Monomorium MG04
Solenopsidini

Monomorium madecassum
Monomorium pharaonis
Monomorium hanneli
Adelomyrmex SC01
Cryptomyrmex boltoni
Baracidris sitra
Adelomyrmex paratristani
Monomorium ferodens
Monomorium fisheri
Monomorium sechellense
Anillomyrma decamera
Anillomyrma AFRC TZ 01
Myrmicaria carinata
Myrmicaria exigua
Monomorium brocha
Monomorium latinode
Tyrannomyrmex legatus
Solenopsis molesta
Solenopsis succinea
Solenopsis xyloni
Solenopsis mameti
Solenopsis fugax
Solenopsis ZA02
Solenopsis papuana
Carebarella PA01
Kempfidris inusualis
Tropidomyrmex elianae
Austromorium flavigaster
Monomorium antarcticum
Monomorium denticulatum
Oxyepoecus vezenyii
Megalomyrmex silvestrii
Rogeria stigmatica
Bariamyrma hispidula
Dolopomyrmex pilatus
Rogeria nevadensis
Stegomyrmex manni
Messor denticornis
Messor wasmanni
Aphaenogaster occidentalis
Goniomma blanci
Oxyopomyrmex santschii

Stenammini Aphaenogaster araneoides


Aphaenogaster pythia
Aphaenogaster swammerdami
Stenamma striatulum
Stenamma dyscheres
Stenamma expolitum
Aphaenogaster albisetosa
Messor andrei
Pogonomyrmex subdentatus

Pogonomyrmecini Pogonomyrmex vermiculatus


Pogonomyrmex imberbiculus
Pogonomyrmex striatinodus
Hylomyrma blandiens
Pogonomyrmex angustus

Myrmicini
Myrmica striolagaster
Myrmica tahoensis
Myrmica rugiventris
Myrmica kotokui
Manica bradleyi
Manica rubida

100 75 50 25 0
million years ago
Fig. 3. (a, b) Biogeographical history of Myrmicinae as inferred from an analysis with lagrange, using a dec (dispersal-extinction-cladogenesis)
model. The ancestral range inheritance scenario with the highest relative probability is indicated for the two descending branches of each node. The
lower half of each sphere pertains to the lower descending branch; the upper half to the upper descending branch. Most (60%) of the relative probabilities
are greater than 0.70, but for some nodes the most probable scenario has weak support and there are alternative scenarios of only slightly lower relative
probability. See Table S7 for a more detailed account of range inheritance scenarios for the major lineages.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


Phylogeny and evolution of myrmicine ants 75

Cardiocondyla MY01
Cardiocondyla thoracica
Cardiocondyla mauritanica
Ocymyrmex fortior cf
Cataulacus ebrardi
Cataulacus hispidulus
Atopomyrmex mocquerysi
Nesomyrmex MG08
Nesomyrmex MG01
Nesomyrmex madecassus
Nesomyrmex echinatinodis

Nearctic Nesomyrmex wilda


Xenomyrmex floridanus
Carebara alperti
Carebara AU01
Neotropical Carebara affinis
Carebara urichi
Carebara nosindambo

Australasian Carebara vidua


Diplomorium longipenne
Vitsika suspicax
Myrmisaraka brevis
Indomalayan Malagidris jugum
Malagidris belti
Royidris notorthotenes
Afrotropical Royidris longiseta
Eutetramorium mocquerysi
Metapone madagascarica

Palearctic Metapone PG01


Liomyrmex gestroi
Podomyrma silvicola
Huberia brounii
Monomorium criniceps
Monomorium destructor
Monomorium robustior
Dacetinops ignotus cf
Crematogaster emeryana
Crematogaster kelleri
Crematogaster modiglianii
Crematogaster acuta
Crematogaster sordidula
Meranoplus castaneus
Meranoplus radamae
Dilobocondyla borneensis cf
Secostruma MY01
Pristomyrmex orbiceps
Pristomyrmex punctatus
Perissomyrmex snyderi
Recurvidris TH01
Adlerzia froggatti
Lophomyrmex ambiguus
Lordomyrma bhutanensis cf
Lordomyrma desupra
Ancyridris polyrhachioides nr
Crematogastrini Propodilobus pingorum
Acanthomyrmex ferox
Mayriella ebbei
Myrmecina americana
Myrmecina graminicola
Myrmecina PG01
Dacatria templaris
Proatta butteli
Tetramorium severini
Tetramorium spinosum
Tetramorium MG125
Rhoptromyrmex transversinodis
Anergates atratulus
Teleutomyrmex kutteri
Strongylognathus testaceus
Tetramorium caespitum cf
Tetramorium venator
Tetramorium validiusculum
Tetramorium CF01
Tetramorium taylori
Myrmicine genus26 PH02
Vollenhovia emeryi
Calyptomyrmex kaurus
Cyphoidris exalta
Dicroaspis KM01
Kartidris sparsipila
Terataner MG02
Terataner MG27
Terataner bottegoi
Melissotarsus insularis
Rhopalomastix rothneyi
Chalepoxenus kutteri
Myrmoxenus bernardi
Temnothorax rugatulus
Temnothorax poeyi

other Myrmicinae Leptothorax muscorum complex


Formicoxenus diversipilosus
Harpagoxenus sublaevis

Fig 3a Vombisidris bilongrudi


Gauromyrmex acanthinus
Rotastruma recava
Rotastruma stenoceps
Romblonella scrobifera
Poecilomyrma myrmecodiae
Paratopula TH01
Tetheamyrma subspongia

100 75 50 25 0
million years ago

Fig. 3. Continued.

The genus Epelysidris stat.r. is resurrected from synonymy tauricus, tramieri and zabelini. Two of these new combinations
under Monomorium to contain the single known species become secondary junior homonyms, T. spinosus (Arnoldi) and
E. brocha (comb.r.). Its putative sister species ‘Monomorium’ T. tauricus (Radchenko), which are here replaced (nom.n.) with
latinode is very different morphologically and a new genus T. sentosus and T. inquilinus, respectively.
will be erected for it in a forthcoming publication (B. Bolton, Myrmoxenus is synonymized (syn.n.) under Temnothorax,
personal communication). generating the following new combinations in Temnothorax:
We support the synonymy of Carebarella under Solenop- adlerzi, africana, algeriana, bernardi, birgitae, corsica, gor-
sis, generating the following new combinations in Solenopsis: diagini, kraussei, ravouxi, stumperi, tamarae, and zaleskyi. Two
alvarengai, condei. The type species of Carebarella (bicolor) of these new combinations, T. kraussei (Emery) and T. tama-
was already combined with Solenopsis by Pacheco & Mackay rae (Arnoldi), create secondary junior homonyms among two
(2013). Although they declined to propose new generic syn- species already in Temnothorax, T. kraussei (Emery) and T.
onymy, their new combination effectively did this, although it tamarae (Radchenko), which are here replaced (nom. n.) with
left the other two names orphaned. T. mediterraneus and T. georgicus, respectively.
Anisopheidole is synonymized (syn.n.) under Pheidole, gen- Protomognathus is synonymized (syn. n.) under Temnothorax,
erating the following revived combination in Pheidole: antipo- generating the following new combination in Temnothorax:
dum. americanus.
Machomyrma is synonymized (syn.n.) under Pheidole, gener- Rhoptromyrmex is synonymized (syn.n.) under Tetramorium,
ating the following new combination in Pheidole: dispar. generating the following new combinations (unless otherwise
Chalepoxenus is synonymized (syn.n.) under Temnothorax, noted) in Tetramorium: caritum, critchleyi, globulinodis, mayri,
generating the following new combinations in Temnothorax: melleum (comb.r.), opacum, rawlinsoni, schmitzi, transversin-
brunneus, hyrcanus, kutteri, muellerianus, spinosus, tarbinskii, odis and wroughtonii. One of these, T. mayri Forel, creates a

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


76 P. S. Ward et al.

secondary junior homonym of a species already in Tetramor- requires further study. Sixteen extant genera that we did not
ium, T. mayri (Mann), which is here replaced (nom.n.) with sequence in this study are indicated with an asterisk (*); they
T. solomonensis. The new combination T. transversinodis have been placed provisionally in tribes on the basis of close
(Mayr) is a secondary senior homonym of an invalid name, morphological or genetic similarity to sequenced taxa. We
T. transversinodis (Enzmann), which therefore does not require cite the author and year of publication of each tribe name;
a replacement name. equivalent information for genus names is available in AntCat
Anergates is synonymized (syn.n.) under Tetramorium, gen- (http://antcat.org/).
erating the following new combination in Tetramorium: atrat-
ulum. Tribe Myrmicini Lepeletier de Saint-Fargeau 1835
Teleutomyrmex is synonymized (syn.n.) under Tetramorium, Constituent genera. Manica, Myrmica, †Plesiomyrmex and
generating the following new combinations in Tetramorium: †Protomyrmica.
kutteri, schneideri. Tetramorium schneideri (Kutter) becomes a
secondary junior homonym, here replaced with T. inquilinum
(nom.n.). Note. Eutetramorium, Huberia, Hylomyrma, Pogonomyrmex
Strongylognathus Mayr is also a synonym of Tetramorium and Secostruma are excluded from Myrmicini.
Mayr, but it has priority over Tetramorium. Transfer of the hun-
dreds of valid Tetramorium species names to Strongylognathus, Tribe Pogonomyrmecini trib.n.
with consequent changes in spelling to many of them, would http://zoobank.org/urn:lsid:zoobank.org:act:BD404053-2C51-
create considerable confusion. In this instance a ruling by the 412B-9291-4A7E9D323A95.
International Commission on Zoological Nomenclature would
be required to validate a reversal of precedence (ICZN Article
Type genus. Pogonomyrmex
23.9.3). In the interim we maintain current usage and refrain
from synonymizing the two genera.
Trichomyrmex stat.r. is resurrected from synonymy under Worker diagnosis. Myrmicine ants of moderate to large
Monomorium, to contain the Monomorium species in the scabri- size (head width 0.73–2.90 mm, head length 0.78–2.50 mm);
ceps- and destructor-groups (sensu Bolton, 1987; see also mandible elongate to subtriangular, with 5–8 teeth on mastica-
Radchenko, 1997). Holcomyrmex and Parholcomyrmex are tory margin; psammophore present or absent; antenna with 12
removed from synonymy under Monomorium, and newly syn- segments, the last 4–5 segments forming a weakly expanded
onymized (syn.n.) under Trichomyrmex. These actions generate club; palp formula (number of maxillary palp segments, fol-
the following new combinations in Trichomyrmex: aberrans, lowed by number of labial palp segments) 5,4 or 4,3; meso-
abyssinicus, chobauti, chobauti ajjer, criniceps, destructor, soma lacking distinct promesonotal suture or metanotal groove;
emeryi, emeryi laevior, epinotale, glaber, lameerei, mayri, propodeum usually with a pair of spines, but lacking in some
muticus, oscaris, perplexus, robustior, santschii (= santschi- species; propodeal lobes usually prominent, and rounded or
anum, unnecessary replacement name) and scabriceps; and spiniform; metasternum with a deep V-shaped or U-shaped
the following revived combinations in Trichomyrmex: rogeri median emargination running forward from the posterior bor-
and wroughtoni (= wroughtonianum, unnecessary replacement der and terminating at or near the metasternal pit, well in front
name). of the metacoxal cavities (Bolton, 1994); metasternal process
large and comprising two prominent teeth; mesotibia and meta-
tibia each with a single apical spur, usually finely barbulate or
Tribal classification pectinate; petiole elongate, with prominent anterior peduncle;
sting well developed and functional; integument sculpture usu-
We propose the recognition of six tribes within Myrmici- ally conspicuous on head, mesosoma, petiole, and postpetiole,
nae. These correspond to the six major (and mutually exclu- typically striate, costate, or rugulose; striae often extending onto
sive) clades that we recover with overwhelming support in all fourth abdominal tergum (first tergum of gaster).
phylogenetic analyses. This represents a considerable reduc-
tion from the 25 tribes currently recognized, but most of those Constituent genera. Hylomyrma and Pogonomyrmex.
tribes are nonmonophyletic or – more often – their recognition
leaves behind a residue of nonmonophyletic taxa. It is this lat-
ter problem that is often overlooked. In the same way that Note. As currently circumscribed Pogonomyrmex is non-
monophyletic ‘satellite genera’ render the genera within which monophyletic, but this can be rectified by assignment of a
they arise paraphyletic (see above), the recognition of small new genus name to the ‘Pogonomyrmex’ angustus-complex. We
well-defined monophyletic tribes, such as Adelomyrmecini, refrain from this action because a detailed phylogenetic analy-
Basicerotini, Lenomyrmecini, Melissotarsini and Myrmicariini, sis of Pogonomyrmecini is currently underway (C. Moreau and
similarly ignores the rump of nonmonophyletic tribes in which R.A. Johnson, personal communication)
these more derived taxa evolved.
In the scheme below, genera known only from fossils are Tribe Stenammini Ashmead 1905
signified by a dagger (†). In many instances their placement = Aphaenogastrini Enzmann 1947 syn.n.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


Phylogeny and evolution of myrmicine ants 77

Constituent genera. Aphaenogaster, Goniomma, Oxy- Tribe Crematogastrini Forel 1893


opomyrmex, Messor, †Paraphaenogaster, Stenamma and = Formicoxenini Forel 1893 (= Cardiocondylini Emery
Veromessor. 1914 = Leptothoracini Emery 1914 = Stereomyrmecini
Emery 1914 = Podomyrmini Emery 1924 = Solenomyrmini
Donisthorpe 1943) syn.n.
Note. Veromessor is resurrected from synonymy (above).
= Cataulacini Emery 1895 syn.n.
The reclassification of ‘Aphaenogaster’ awaits more detailed
= Tetramoriini Emery 1895 (= Anergatini Emery 1914
phylogenetic analysis of lineage diversity within Stenammini.
= Teleutomyrmecini Kutter 1950) syn.n.
= Melissotarsini Emery 1901 syn.n.
Tribe Solenopsidini Forel 1893 = Myrmecinini Ashmead 1905 syn.n.
= Myrmicariini Forel 1893 syn.n. = Metaponini Forel 1911 syn.n.
= Monomoriini Emery 1914 = Lophomyrmecini Emery 1914 syn.n.
= Megalomyrmecini Dlussky & Fedoseeva 1988 = Meranoplini Emery 1914 syn.n.
= Stegomyrmecini Wheeler 1922 syn.n. = Ocymyrmecini Emery 1914 syn.n.
= Adelomyrmecini Fernández 2004 syn.n. = Pheidologetonini Emery 1914 syn.n.
= Proattini Forel 1917 syn.n.
= Hypopomyrmecina Brown 1952 syn.n.
Constituent genera. Adelomyrmex, Anillomyrma, Austromo- = Calyptomyrmecini Dlussky & Fedoseeva 1988 syn.n.
rium, Baracidris, Bariamyrma, *Bondroitia, Cryptomyrmex, = Liomyrmecini Bolton 2003 syn.n.
Dolopomyrmex, Epelysidris, Kempfidris, Megalomyrmex, = Paratopulini Bolton 2003 syn.n.
Monomorium, Myrmicaria, Oxyepoecus, Rogeria, Solenopsis,
Stegomyrmex, Syllophopsis, Tropidomyrmex and Tyranno-
myrmex. Constituent genera. Acanthomyrmex, Adlerzia, Ancyridris,
Atopomyrmex, Calyptomyrmex, Cardiocondyla, Carebara,
Cataulacus, Crematogaster, Cyphoidris, Dacatria, Dacetinops,
Note. Carebarella is considered a synonym of Solenopsis Dicroaspis, Dilobocondyla, Diplomorium, †Enneamerus,
(above). Monomorium remains a heterogeneous, nonmono- †Eocenomyrma, Eutetramorium, Formicoxenus, *For-
phyletic group. Extant genera excluded from Solenopsidini mosimyrma, Gauromyrmex, *Goaligongidris, Harpago-
include Adlerzia, Allomerus, Carebara, Diplomorium, xenus, Huberia, †Hypopomyrmex, *Indomyrma, Kartidris,
Mayriella and Tranopelta. *Lasiomyrma, Leptothorax, Liomyrmex, †Lonchomyrmex,
Lophomyrmex, Lordomyrma, Malagidris, Mayriella, Melis-
sotarsus, Meranoplus, Metapone, Myrmecina, Myrmisaraka,
Tribe Attini F. Smith 1858
Nesomyrmex, Ocymyrmex, †Oxyidris, †Parameranoplus,
= Pheidolini Emery 1877 (= Anergatidini Emery 1922) syn.n.
Paratopula, Perissomyrmex, *Peronomyrmex, Podomyrma, Poe-
= Dacetini Forel 1892 (= Epopostrumina Brown 1952
cilomyrma, Pristomyrmex, Proatta, Propodilobus, Recurvidris,
= Orectognathina Brown 1952 = Strumigenina Brown 1952)
Rhopalomastix, Romblonella, *Rostromyrmex, Rotastruma,
syn. n.
Royidris, Secostruma, *Stereomyrmex, †Stigmomyrmex,
= Ochetomyrmecini Emery 1914 syn.n.
†Stiphromyrmex, Strongylognathus, Temnothorax, Terataner,
= Cephalotini M. R. Smith 1949 (= Cryptoceridae F. Smith
Tetheamyrma, Tetramorium, Trichomyrmex, Vitsika, Vollen-
1853) syn.n.
hovia, Vombisidris and Xenomyrmex.
= Basicerotini Brown 1949 syn.n.
= Phalacromyrmecini Dlussky & Fedoseeva 1988 syn.n.
= Blepharidattini Wheeler & Wheeler 1991 syn.n.
= Lenomyrmecini Bolton 2003 syn.n. Fossil taxa unplaced to tribe

†Afromyrma, †Attopsis, †Bilobomyrma, †Boltonidris,


Constituent genera. Acanthognathus, Acromyrmex, †Brachytarsites, †Cephalomyrmex, †Clavipetiola,
Allomerus, Apterostigma, Atta, †Attaichnus, Basiceros, †Electromyrmex, †Eocenidris, †Eomyrmex, †Fallomyrma,
Blepharidatta, Cephalotes, *Chimaeridris, Colobostruma, †Fushunomyrmex, †Ilemomyrmex, †Miosolenopsis,
*Cyatta, Cyphomyrmex, Daceton, Diaphoromyrma, †Myrmecites, †Orbigastrula, †Quadrulicapito,
Epopostruma, Eurhopalothrix, *Ishakidris, Kalathomyrmex, †Quineangulicapito, †Sinomyrmex, †Solenopsites,
Lachnomyrmex, Lenomyrmex, Mesostruma, Microdaceton, †Sphaerogasterites, †Wumyrmex and †Zhangidris.
*Mycetagroicus, Mycetarotes, *Mycetophylax, Mycetosoritis,
Mycocepurus, Myrmicocrypta, Ochetomyrmex, Octostruma,
Orectognathus, *Paramycetophylax, Phalacromyrmex, Phei- Excluded from Myrmicinae
dole, Pilotrochus, Procryptocerus, Protalaridris, *Pseudoatta,
Rhopalothrix, *Sericomyrmex, Strumigenys, Talaridris, Tra- The tribe Ankylomyrmini Bolton and its sole genus, Anky-
chymyrmex, Tranopelta and Wasmannia. lomyrma Bolton, are hereby transferred to the subfamily

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81


78 P. S. Ward et al.

Agroecomyrmecinae, in accordance with our finding that Table S5. Species distribution matrix, based on six biogeo-
Ankylomyrma is sister to Tatuidris (Fig. 1). graphic areas: Neotropical (T), Nearctic (N), Palearctic (P),
Afrotropical (E), Indomalayan (O) and Australasian regions
(A).
Concluding remarks Table S6. Support values from various modes of anal-
ysis for selected clades of interest. These include
With respect to taxonomic diversity, geographical distribution
and ecological breadth, myrmicine ants are the most successful maximum-likelihood analysis of the full dataset, Bayesian
of all ant subfamilies. Our results indicate that the diversifica- analysis of the full dataset and treatments designed to
tion of crown-group Myrmicinae began about 100 Ma, and was address base frequency heterogeneity.
initially concentrated in the New World tropics. Repeated dis- Table S7. Range inheritance inferences for major clades of
persal to, and radiation in, other biogeographic regions was a Myrmicinae, based on lagrange analysis.
key feature of myrmicine evolution. The subfamily now com-
prises more than 6000 described species – the actual number of
species could well be twice this – and occupies all major terres- Acknowledgements
trial ecoregions. Analysis of our 251-taxon 11-gene dataset, in
conjunction with 27 fossil calibration points, reveals the broad We thank Rachelle Adams, Gary Alpert, Bob Anderson,
outlines of the evolutionary history of this group, including Himender Bharti, Lech Borowiec, Marek Borowiec, Beto
the existence of six large, mutually exclusive clades. Neverthe- Brandão, Michael Branstetter, Bui Tuan Viet, Chris Burwell,
less considerable challenges remain in myrmicine systematics, Katsuyuki Eguchi, Xavier Espadaler, Rodrigo Feitosa, John
including resolution of the basal polytomy in the Crematogas- Fellowes, Fernando Fernández, Juergen Gadau, David General,
trini clade, reclassification of nonmonophyletic taxa such as Nihara Gunawardene, Yoshiaki Hashimoto, Peter Hawkes, Jür-
Aphaenogaster and Monomorium, and better incorporation of gen Heinze, Milan Janda, Dan Kjar, John Lattke, Jack Longino,
fossil taxa into the phylogenetic scaffold. Much more attention Andrea Lucky, Dirk Mezger, James Pitts, Shauna Price, Eli Sar-
also needs to be directed towards the neglected species-level tax- nat, Steve Shattuck, Rogério Silva, Jeffrey Sosa-Calvo, Andy
onomy of many myrmicine genera. Suarez, Noel Tawatao, Alberto Tinaut, Darren Ward and Alex
Wild for contributing specimens towards this study. Paul Arm-
strong, Matt Kweskin, Eugenia Okonski, Jamie Pettengill and
Hong Zhao provided technical support. Barry Bolton, Brendon
Supporting Information
Boudinot, Marek Borowiec and an anonymous reviewer gave
helpful comments that improved the manuscript. This study
Additional Supporting Information may be found in the online
was supported by NSF grants EF-0431330, DEB-0743542,
version of this article under the DOI reference:
DEB-0842204, and DEB-0949689.
10.1111/syen.12090
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Ward, P.S., Brady, S.G., Fisher, B.L., & Schultz, T.R. (2014) The evolution of myrmicine ants: phylogeny
and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology

Figure S1. Majority-rule consensus trees from partitioned, concatenated Bayesian analyses of data sets
with “wildcard” taxon exclusions. Fig. S1A is based on 248 taxa, and excludes Acanthomyrmex ferox,
Mayriella ebbei, and Xenomyrmex floridanus; Fig. S1B is based on 243 taxa, and excludes Cardiocondyla
mauritanica, Cardiocondyla MY01, Cardiocondyla thoracica, Liomyrmex gestroi, Melissotarsus insularis,
Metapone madagascarica, Metapone PG01, and Ocymyrmex cf. fortior.
)LJXUH6D248WD[D

100 Apterogyna ZA01


Amblyopone australis
89 Stigmatomma pallipes
88 58 Proceratium stictum
69 Paraponera clavata
100 Platythyrea mocquerysi
100 Ankylomyrma coronacantha
Tatuidris WDWXVLD
100 Tatuidris tatusiaB
Aneuretus simoni
100 Dolichoderus pustulatus
100 100 Tetraponera rufonigra
Myrmecia pyriformis
100 100 Nothomyrmecia macrops
Lasius californicus
100 Myrmelachista flavocotea
Acanthoponera minor
100 100 Rhytidoponera chalybaea
Manica bradleyi
100 Manica rubida
100 100 Myrmica kotokui
100 Myrmica rugiventris
Myrmica striolagaster
90 Myrmica tahoensis
100 Pogonomyrmex angustus
99 Hylomyrma blandiens
100 100 Pogonomyrmex striatinodus
100 Pogonomyrmex imberbiculus
Pogonomyrmex subdentatus
100 100 Pogonomyrmex vermiculatus
Aphaenogaster albisetosa
100 Messor andrei
100 100 Stenamma expolitum
100 Stenamma dyscheres
100 Stenamma striatulum
Aphaenogaster pythia
100 Aphaenogaster swammerdami
75 100 Aphaenogaster araneoides
100 Goniomma blanci
100 Oxyopomyrmex santschii
100 Aphaenogaster occidentalis
Messor denticornis
97 Messor wasmanni
99 Stegomyrmex manni
Dolopomyrmex pilatus
100 100 Rogeria nevadensis
Bariamyrma hispidula
100 100 Rogeria stigmatica
85 Austromorium flavigaster
100 Monomorium antarcticum
Monomorium denticulatum
99 100 Oxyepoecus vezenyii
Kempfidris inusualis
100 100 Tropidomyrmex elianae
Carebarella PA01
100 100 Solenopsis papuana
100 100 Solenopsis fugax
100 Solenopsis ZA02
100 Solenopsis mameti
100 100 Solenopsis xyloni
Solenopsis molesta
Solenopsis succinea
100 Megalomyrmex silvestrii
72 79 Tyrannomyrmex legatus
Monomorium brocha
72 Monomorium latinode
100 100 Monomorium sechellense
Monomorium ferodens
87 Monomorium fisheri
100 Adelomyrmex paratristani
100 Baracidris sitra
100 Adelomyrmex SC01
100 Cryptomyrmex boltoni
94 Anillomyrma AFRC TZ 01
100 Anillomyrma decamera
Myrmicaria carinata
Myrmicaria exigua
89 Monomorium hanneli
100 Monomorium pharaonis
55 100 Monomorium madecassum
100 Monomorium ergatogyna
100 Monomorium fieldi nr
Monomorium exiguum
100 Monomorium MG04
100 Ochetomyrmex neopolitus
100 Ochetomyrmex semipolitus
Tranopelta subterranea
100 Tranopelta gilva
92 Diaphoromyrma sofiae
89 100 Lachnomyrmex scrobiculatus
100 Blepharidatta conops
Allomerus octoarticulatus
100 Wasmannia auropunctata
95 72 Acanthognathus ocellatus
Daceton armigerum
95 Lenomyrmex colwelli
100 Microdaceton tibialis
100 100 Orectognathus versicolor
94 100 100 Colobostruma unicorna
Epopostruma monstrosa
100 Mesostruma browni
100 Apterostigma auriculatum
Mycocepurus goeldii
100 Myrmicocrypta infuscata cf
100 59 Kalathomyrmex emeryi
Mycetarotes acutus
100 Mycetosoritis hartmanni
100 Cyphomyrmex cornutus
93 Cyphomyrmex longiscapus cf
98 100 Acromyrmex versicolor
Atta texana
100 Trachymyrmex arizonensis
99 100 Procryptocerus scabriusculus
Cephalotes atratus
Cephalotes texanus
100 Pheidole fimbriata
100 Pheidole rhea
100 Pheidole clydei
69 100 Pheidole hyatti
100 Pheidole lucioccipitalis
96 100 Pheidole pallidula
52 Anisopheidole antipodum
Machomyrma dispar
100 Pheidole longispinosa
Protalaridris armata
100 Rhopalothrix isthmica
90 Basiceros manni
100 100 Octostruma EC01
82 100 100 Eurhopalothrix australis
Eurhopalothrix gravis
100 Talaridris mandibularis
Phalacromyrmex fugax
100 Pilotrochus besmerus
100 6WUXPLJHQ\V ambatrix
100 100 6WUXPLJHQ\V nitens
6WUXPLJHQ\V gundlachi
100 100 Strumigenys biolleyi
6WUXPLJHQ\V erynnes
100 100 6WUXPLJHQ\V ludovici
100 Strumigenys rogeri
87 Strumigenys coveri
Strumigenys dicomas
100 6WUXPLJHQ\VPD[LOODULV
95 6WUXPLJHQ\V olsoni
96 97 Strumigenys exiguaevitae
100 6WUXPLJHQ\V membranifera
91 6WUXPLJHQ\V ocypete
100 6WUXPLJHQ\V chiricahua
100 Strumigenys emmae
Strumigenys chyzeri
Strumigenys godeffroyi
Tetheamyrma subspongia
100 Gauromyrmex acanthinus
69 100 Vombisidris bilongrudi
74 99 Harpagoxenus sublaevis
Formicoxenus diversipilosus
100 Leptothorax muscorum complex
75 Temnothorax poeyi
89 100 Temnothorax rugatulus
Chalepoxenus kutteri
100 Myrmoxenus bernardi
Melissotarsus insularis
100 Rhopalomastix rothneyi
100 100 Terataner bottegoi
100 Terataner MG02
Terataner MG27
100 98 100 Kartidris sparsipila
Cyphoidris exalta
100 Dicroaspis KM01
100 100 Calyptomyrmex kaurus
Myrmicine genus26 PH02
100 100 Vollenhovia emeryi
7HWUDPRULXPYHQDWRU
100 Tetramorium validiusculum
Tetramorium CF01
100 100 Tetramorium taylori
100 Tetramorium caespitum cf
100 Strongylognathus testaceus
Anergates atratulus
100 Teleutomyrmex kutteri
97 Rhoptromyrmex transversinodis
100 Tetramorium MG125
Tetramorium severini
87 Tetramorium spinosum
Dacetinops ignotus cf
71 Lophomyrmex ambiguus
Adlerzia froggatti
100 Recurvidris TH01
Dacatria templaris
100 Proatta butteli
Dilobocondyla borneensis cf
100 Secostruma MY01
100 Myrmecina PG01
Myrmecina americana
100 Myrmecina graminicola
100 Perissomyrmex snyderi
Pristomyrmex orbiceps
100 Pristomyrmex punctatus
54 Propodilobus pingorum
94 Ancyridris polyrhachioides nr
Lordomyrma bhutanensis cf
Lordomyrma desupra
100 Paratopula TH01
100 Poecilomyrma myrmecodiae
100 Romblonella scrobifera
Rotastruma recava
Rotastruma stenoceps
100 Diplomorium longipenne
100 Carebara vidua
66 100 100 Carebara nosindambo
100 Carebara alperti
100 Carebara AU01
Carebara urichi
100 &DUHEDUDDIILQLV
Meranoplus castaneus
83 100 Meranoplus radamae
100 Crematogaster acuta
100 Crematogaster sordidula
100 Crematogaster modiglianii
Crematogaster adrepens
100 Crematogaster emeryana
100 Nesomyrmex echinatinodis
100 Nesomyrmex wilda
94 Nesomyrmex madecassus
90 Nesomyrmex MG01
Nesomyrmex MG08
75 100 Atopomyrmex mocquerysi
Cataulacus ebrardi
98 Cataulacus hispidulus
100 Ocymyrmex fortior cf
100 Cardiocondyla mauritanica
Cardiocondyla MY01
100 Cardiocondyla thoracica
100 Monomorium robustior
Monomorium criniceps
100 Monomorium destructor
100 Huberia brounii
Podomyrma silvicola
100 100 Liomyrmex gestroi
100 Metapone madagascarica
Metapone PG01
100 100 Eutetramorium mocquerysi
Royidris notorthotenes
82 100 Royidris longiseta
100 0DODJLGULVMXJXP
100 MDODJLGULVEHOWL
Myrmisaraka brevis
Vitsika suspicax
)LJXUH6EWD[D
100 Apterogyna ZA01
Amblyopone australis
84 Stigmatomma pallipes
83 63 Proceratium stictum
70 Paraponera clavata
Platythyrea mocquerysi
100 Ankylomyrma coronacantha
100 Tatuidris WDWXVLD
100 Tatuidris tatusiaB
100 Aneuretus simoni
Dolichoderus pustulatus
100 Tetraponera rufonigra
100 Myrmecia pyriformis
100 Nothomyrmecia macrops
100 Lasius californicus
100 Myrmelachista flavocotea
Acanthoponera minor
100 Rhytidoponera chalybaea
100 100 Manica bradleyi
Manica rubida
100 Myrmica kotokui
100 Myrmica rugiventris
100 Myrmica striolagaster
92 Myrmica tahoensis
100 Pogonomyrmex angustus
99 Hylomyrma blandiens
100 Pogonomyrmex striatinodus
100 Pogonomyrmex imberbiculus
100 Pogonomyrmex subdentatus
100 Pogonomyrmex vermiculatus
100 100 Aphaenogaster albisetosa
Messor andrei
100 Stenamma expolitum
100
100 Stenamma dyscheres
100 Stenamma striatulum
Aphaenogaster pythia
100 Aphaenogaster swammerdami
75 100 Aphaenogaster araneoides
100 Goniomma blanci
Oxyopomyrmex santschii
100 Aphaenogaster occidentalis
100 Messor denticornis
97 Messor wasmanni
99 Stegomyrmex manni
100 Dolopomyrmex pilatus
100 Rogeria nevadensis
Bariamyrma hispidula
100 100 Rogeria stigmatica
82 Austromorium flavigaster
100 Monomorium antarcticum
Monomorium denticulatum
99 100 Oxyepoecus vezenyii
.HPSILGULVLQXVXDOLV
100 100 Tropidomyrmex elianae
Carebarella PA01
100 100 Solenopsis papuana
100 100 Solenopsis fugax
Solenopsis ZA02
100 Solenopsis mameti
100 Solenopsis xyloni
100 100 Solenopsis molesta
Solenopsis succinea
Megalomyrmex silvestrii
100 Tyrannomyrmex legatus
66 75 Monomorium brocha
Monomorium latinode
57 Monomorium sechellense
100 100 Monomorium ferodens
77 Monomorium fisheri
100 Adelomyrmex paratristani
100 Baracidris sitra
100 Adelomyrmex SC01
100 Cryptomyrmex boltoni
94 Anillomyrma AFRC TZ 01
100 Anillomyrma decamera
Myrmicaria carinata
Myrmicaria exigua
87 Monomorium hanneli
100 Monomorium pharaonis
56 100 Monomorium madecassum
100 Monomorium ergatogyna
100 Monomorium fieldi nr
Monomorium exiguum
100 Monomorium MG04
100 Ochetomyrmex neopolitus
100 Ochetomyrmex semipolitus
100 Tranopelta subterranea
Tranopelta gilva
90 Diaphoromyrma sofiae
89 Lachnomyrmex scrobiculatus
100 Blepharidatta conops
100 Allomerus octoarticulatus
100 Wasmannia auropunctata
95 71 Acanthognathus ocellatus
92 Daceton armigerum
Lenomyrmex colwelli
100 Microdaceton tibialis
100 Orectognathus versicolor
100 100 Colobostruma unicorna
92 100 Epopostruma monstrosa
Mesostruma browni
100 Apterostigma auriculatum
100 Mycocepurus goeldii
100 Myrmicocrypta infuscata cf
100 66 Kalathomyrmex emeryi
Mycetarotes acutus
100 Mycetosoritis hartmanni
100 Cyphomyrmex cornutus
92 Cyphomyrmex longiscapus cf
98 100 Acromyrmex versicolor
Atta texana
Trachymyrmex arizonensis
100 Procryptocerus scabriusculus
100
99 Cephalotes atratus
Cephalotes texanus
100 Pheidole fimbriata
100 Pheidole rhea
100 Pheidole clydei
67 100 Pheidole hyatti
100 Pheidole lucioccipitalis
Pheidole pallidula
95 100 Anisopheidole antipodum
52
Machomyrma dispar
100 Pheidole longispinosa
100 Protalaridris armata
Rhopalothrix isthmica
91 Basiceros manni
100 Octostruma EC01
100 Eurhopalothrix australis
81 100 100
Eurhopalothrix gravis
100 Talaridris mandibularis
Phalacromyrmex fugax
100 Pilotrochus besmerus
6WUXPLJHQ\V ambatrix
100 100 6WUXPLJHQ\V nitens
100
6WUXPLJHQ\V gundlachi
100 100 Strumigenys biolleyi
6WUXPLJHQ\V erynnes
100 6WUXPLJHQ\V ludovici
100 Strumigenys rogeri
100
87 Strumigenys coveri
Strumigenys dicomas
100 6WUXPLJHQ\VPD[LOODULV
95 6WUXPLJHQ\V olsoni
96 98 Strumigenys exiguaevitae
100 6WUXPLJHQ\V membranifera
6WUXPLJHQ\V ocypete
92 6WUXPLJHQ\V chiricahua
100 Strumigenys emmae
100
Strumigenys chyzeri
Strumigenys godeffroyi
Tetheamyrma subspongia
100 Paratopula TH01
100 Poecilomyrma myrmecodiae
59 Romblonella scrobifera
60 100
Rotastruma recava
Rotastruma stenoceps
100 Gauromyrmex acanthinus
78 100 Vombisidris bilongrudi
99 Harpagoxenus sublaevis
82 Formicoxenus diversipilosus
56 Leptothorax muscorum complex
100 Temnothorax poeyi
73 Temnothorax rugatulus
100
Chalepoxenus kutteri
100 100 Myrmoxenus bernardi
Rhopalomastix rothneyi
100 Terataner bottegoi
100
100 Terataner MG02
Terataner MG27
100 Kartidris sparsipila
99 Cyphoidris exalta
Dicroaspis KM01
100 100 Calyptomyrmex kaurus
100
Myrmicine genus26 PH02
100 Vollenhovia emeryi
100 7HWUDPRULXPYHQDWRU
100 Tetramorium validiusculum
Tetramorium CF01
100 Tetramorium taylori
100 Tetramorium caespitum cf
86 100 Strongylognathus testaceus
100
Anergates atratulus
Teleutomyrmex kutteri
100 Rhoptromyrmex transversinodis
97 Tetramorium MG125
100
Tetramorium severini
100 Tetramorium spinosum
Dacatria templaris
100 Proatta butteli
100 Myrmecina PG01
Myrmecina americana
78 Myrmecina graminicola
87 Acanthomyrmex ferox
Mayriella ebbei
100 Propodilobus pingorum
52 Ancyridris polyrhachioides nr
96
Lordomyrma bhutanensis cf
74 Lordomyrma desupra
65 93 Lophomyrmex ambiguus
Adlerzia froggatti
100 Recurvidris TH01
57 Dilobocondyla borneensis cf
100 Secostruma MY01
100 Perissomyrmex snyderi
Pristomyrmex orbiceps
Pristomyrmex punctatus
100 Diplomorium longipenne
100 Carebara vidua
100 100 Carebara nosindambo
100 Carebara alperti
100 Carebara AU01
52 Carebara urichi
100 &DUHEDUDDIILQLV
Meranoplus castaneus
91 100 Meranoplus radamae
100 Crematogaster acuta
100 Crematogaster sordidula
100 Crematogaster modiglianii
Crematogaster adrepens
Crematogaster emeryana
85 Dacetinops ignotus cf
100 Atopomyrmex mocquerysi
99 Cataulacus ebrardi
Cataulacus hispidulus
79 100 Xenomyrmex floridanus
76 100 Nesomyrmex echinatinodis
100 Nesomyrmex wilda
94 Nesomyrmex madecassus
Nesomyrmex MG01
100 Nesomyrmex MG08
100 Monomorium robustior
Monomorium criniceps
100 100 Monomorium destructor
Huberia brounii
100 Podomyrma silvicola
100 100 Eutetramorium mocquerysi
Royidris notorthotenes
89 100 Royidris longiseta
100 MDODJLGULVMXJXP
100 MDODJLGULVEHOWL
Myrmisaraka brevis
Vitsika suspicax
Ward, P.S., Brady, S.G., Fisher, B.L., & Schultz, T.R. (2014) The evolution of myrmicine ants: phylogeny
and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology

Figure S2. Majority-rule consensus trees resulting from Bayesian analyses of single-gene fragments (18S,
28S, Abd-A, EF1aF1, LW Rh, ArgK, Top1, Ubx, Wg, EF1aF2, and CAD) under the partitions and models
described in Table S3b.
)LJXUH618S

Apterogyna ZA01
Platythyrea mocquerysi
78 Paraponera clavata
Amblyopone australis
Stigmatomma pallipes
99 99 Proceratium stictum
Ankylomyrma coronacantha
Tatuidris WDWXVLD
97 Tatuidris tatusiaB
Myrmecia pyriformis
98 80 Nothomyrmecia macrops
96 100 Tetraponera rufonigra
Aneuretus simoni
Dolichoderus pustulatus
Acanthoponera minor
80 Rhytidoponera chalybaea
81 Lasius californicus
Myrmelachista flavocotea
Hylomyrma blandiens
Manica bradleyi
Manica rubida
Pogonomyrmex angustus
91 Pogonomyrmex striatinodus
Pogonomyrmex imberbiculus
Pogonomyrmex subdentatus
Pogonomyrmex vermiculatus
100 58 Myrmica kotokui
Myrmica rugiventris
Myrmica striolagaster
Myrmica tahoensis
Acanthomyrmex ferox
Acromyrmex versicolor
Adelomyrmex paratristani
Adlerzia froggatti
Allomerus octoarticulatus
Anergates atratulus
Anisopheidole antipodum
Aphaenogaster albisetosa
Aphaenogaster araneoides
Aphaenogaster occidentalis
Aphaenogaster pythia
Aphaenogaster swammerdami
Apterostigma auriculatum
58 Atopomyrmex mocquerysi
Atta texana
Austromorium flavigaster
Basiceros manni
Blepharidatta conops
Calyptomyrmex kaurus
Cataulacus ebrardi
Cataulacus hispidulus
Cephalotes atratus
Cephalotes texanus
Chalepoxenus kutteri
Crematogaster emeryana
Cryptomyrmex boltoni
Cyphoidris exalta
Cyphomyrmex cornutus
Cyphomyrmex longiscapus cf
Dacatria templaris
Dacetinops ignotus cf
7HWUDPRULXPYHQDWRU
Diaphoromyrma sofiae
Dicroaspis KM01
Dolopomyrmex pilatus
Eutetramorium mocquerysi
Formicoxenus diversipilosus
Gauromyrmex acanthinus
Goniomma blanci
Harpagoxenus sublaevis
Huberia brounii
Kartidris sparsipila
Lachnomyrmex scrobiculatus
Lenomyrmex colwelli
Leptothorax muscorum complex
59 Liomyrmex gestroi
Lophomyrmex ambiguus
Machomyrma dispar
Megalomyrmex silvestrii
Melissotarsus insularis
Messor andrei
Monomorium antarcticum
Monomorium brocha
Monomorium denticulatum
Monomorium fisheri
.HPSILGULVLQXVXDOLV
Mycetarotes acutus
Mycetosoritis hartmanni
Mycocepurus goeldii
Myrmicaria carinata
Myrmicaria exigua
MDODJLGULVMXJXP
MDODJLGULVEHOWL
Myrmisaraka brevis
Vitsika suspicax
Myrmicocrypta infuscata cf
Myrmoxenus bernardi
Nesomyrmex echinatinodis
Nesomyrmex madecassus
Nesomyrmex MG01
Nesomyrmex MG08
Nesomyrmex wilda
Octostruma EC01
Ocymyrmex fortior cf
Paratopula TH01
Perissomyrmex snyderi
Phalacromyrmex fugax
Pheidole clydei
Pheidole fimbriata
Pheidole hyatti
Pheidole longispinosa
Pheidole lucioccipitalis
Pheidole pallidula
Pheidole rhea
Pilotrochus besmerus
Podomyrma silvicola
Poecilomyrma myrmecodiae
Pristomyrmex orbiceps
Pristomyrmex punctatus
Proatta butteli
Procryptocerus scabriusculus
Propodilobus pingorum
Protalaridris armata
Recurvidris TH01
Rhopalomastix rothneyi
Rhopalothrix isthmica
Rhoptromyrmex transversinodis
Rogeria stigmatica
Romblonella scrobifera
Rotastruma recava
Rotastruma stenoceps
Stegomyrmex manni
Stenamma dyscheres
80 Stenamma expolitum
Stenamma striatulum
Strongylognathus testaceus
Teleutomyrmex kutteri
Temnothorax poeyi
Temnothorax rugatulus
Tetheamyrma subspongia
Tetramorium caespitum cf
Tetramorium CF01
Tetramorium MG125
Tetramorium severini
Tetramorium spinosum
Tetramorium taylori
Tetramorium validiusculum
Trachymyrmex arizonensis
Tropidomyrmex elianae
Tyrannomyrmex legatus
Vollenhovia emeryi
Vombisidris bilongrudi
100 Wasmannia auropunctata
Acanthognathus ocellatus
99 Microdaceton tibialis
Anillomyrma AFRC TZ 01
92 Anillomyrma decamera
Bariamyrma hispidula
60 Mayriella ebbei
Daceton armigerum
78 Kalathomyrmex emeryi
Dilobocondyla borneensis cf
51 Secostruma MY01
Meranoplus castaneus
100 Meranoplus radamae
Metapone madagascarica
84 Metapone PG01
Monomorium hanneli
96 Rogeria nevadensis
Royidris notorthotenes
68 Royidris longiseta
Myrmicine genus26 PH02
97 Xenomyrmex floridanus
Ochetomyrmex neopolitus
99 Ochetomyrmex semipolitus
Tranopelta subterranea
99 Tranopelta gilva
Ancyridris polyrhachioides nr
Lordomyrma bhutanensis cf
77 Lordomyrma desupra
Cardiocondyla mauritanica
Cardiocondyla MY01
98 Cardiocondyla thoracica
86 Eurhopalothrix australis
Eurhopalothrix gravis
58 Talaridris mandibularis
100 Oxyopomyrmex santschii
Messor denticornis
100 Messor wasmanni
Monomorium criniceps
Monomorium destructor
100 Monomorium robustior
Terataner bottegoi
Terataner MG02
Terataner MG27
85 Monomorium latinode
86 Monomorium sechellense
Adelomyrmex SC01
Oxyepoecus vezenyii
74 Colobostruma unicorna
Epopostruma monstrosa
Mesostruma browni
Orectognathus versicolor
71 Diplomorium longipenne
55 Carebara vidua
88 97 Carebara nosindambo
Carebara alperti
97 Carebara AU01
Carebara urichi
&DUHEDUDDIILQLV
Crematogaster acuta
90 61 Crematogaster adrepens
Crematogaster modiglianii
68 Crematogaster sordidula
Myrmecina americana
Myrmecina graminicola
Myrmecina PG01
96 Baracidris sitra
Solenopsis fugax
Solenopsis xyloni
Carebarella PA01
87 79 Solenopsis mameti
Solenopsis molesta
Solenopsis papuana
Solenopsis succinea
Solenopsis ZA02
Monomorium exiguum
96 Monomorium madecassum
Monomorium MG04
89 Monomorium pharaonis
Monomorium ergatogyna
Monomorium fieldi nr
6WUXPLJHQ\V ambatrix
6WUXPLJHQ\V erynnes
6WUXPLJHQ\V ludovici
98 Strumigenys biolleyi
6WUXPLJHQ\V gundlachi
98 99 6WUXPLJHQ\V nitens
6WUXPLJHQ\V membranifera
Strumigenys godeffroyi
Monomorium ferodens
Strumigenys chiricahua
Strumigenys exiguaevitae
69 6WUXPLJHQ\VPD[LOODULV
Strumigenys ocypete
Strumigenys olsoni
Strumigenys chyzeri
Strumigenys emmae
100 Strumigenys rogeri
Strumigenys coveri
Strumigenys dicomas
)LJXUH628S

Apterogyna ZA01
Paraponera clavata
98 Proceratium stictum
Amblyopone australis
100 Stigmatomma pallipes
100 Ankylomyrma coronacantha
Tatuidris WDWXVLD
Tatuidris tatusiaB
100 Platythyrea mocquerysi
Acanthoponera minor
51 100 Rhytidoponera chalybaea
Lasius californicus
100 Myrmelachista flavocotea
Myrmecia pyriformis
84 Nothomyrmecia macrops
57 96 Tetraponera rufonigra
Aneuretus simoni
Dolichoderus pustulatus
Acanthognathus ocellatus
88 Adelomyrmex paratristani
Adelomyrmex SC01
Adlerzia froggatti
Ancyridris polyrhachioides nr
Apterostigma auriculatum
Baracidris sitra
Basiceros manni
Calyptomyrmex kaurus
Carebarella PA01
Cephalotes atratus
Cephalotes texanus
Cryptomyrmex boltoni
Cyphoidris exalta
Dacatria templaris
Dacetinops ignotus cf
Diaphoromyrma sofiae
Dicroaspis KM01
Eutetramorium mocquerysi
Gauromyrmex acanthinus
69 Goniomma blanci
Huberia brounii
Kalathomyrmex emeryi
Kartidris sparsipila
Lachnomyrmex scrobiculatus
Lordomyrma bhutanensis cf
Lordomyrma desupra
Mayriella ebbei
Melissotarsus insularis
Messor andrei
Monomorium antarcticum
Monomorium ferodens
Mycocepurus goeldii
Vitsika suspicax
Myrmicine genus26 PH02
Myrmicocrypta infuscata cf
Myrmoxenus bernardi
Octostruma EC01
Oxyepoecus vezenyii
Perissomyrmex snyderi
Pilotrochus besmerus
Podomyrma silvicola
Poecilomyrma myrmecodiae
Proatta butteli
Procryptocerus scabriusculus
Propodilobus pingorum
Protalaridris armata
Recurvidris TH01
Rhopalomastix rothneyi
Rhopalothrix isthmica
Rogeria nevadensis
100 Rogeria stigmatica
Solenopsis papuana
Stenamma dyscheres
Stenamma striatulum
Temnothorax poeyi
Temnothorax rugatulus
Tetheamyrma subspongia
Tyrannomyrmex legatus
Vollenhovia emeryi
Vombisidris bilongrudi
100 Xenomyrmex floridanus
Allomerus octoarticulatus
100 Wasmannia auropunctata
Anillomyrma AFRC TZ 01
54 Anillomyrma decamera
Aphaenogaster albisetosa
100 Lophomyrmex ambiguus
Atopomyrmex mocquerysi
52 Myrmisaraka brevis
Austromorium flavigaster
59 Monomorium brocha
Blepharidatta conops
100 Chalepoxenus kutteri
Cataulacus ebrardi
100 Cataulacus hispidulus
Crematogaster acuta
100 Crematogaster sordidula
Dilobocondyla borneensis cf
91 Secostruma MY01
Megalomyrmex silvestrii
99 Monomorium denticulatum
Meranoplus castaneus
100 Meranoplus radamae
Royidris notorthotenes
100 Royidris longiseta
Myrmicaria carinata
99 Myrmicaria exigua
MDODJLGULVMXJXP
100 MDODJLGULVEHOWL
Nesomyrmex echinatinodis
100 Nesomyrmex wilda
Ochetomyrmex neopolitus
81 Ochetomyrmex semipolitus
Ocymyrmex fortior cf
100 Phalacromyrmex fugax
Tranopelta subterranea
57 Tranopelta gilva
100 Acanthomyrmex ferox
Pristomyrmex orbiceps
98 Pristomyrmex punctatus
58 Cardiocondyla MY01
Cardiocondyla mauritanica
98 Cardiocondyla thoracica
90 Crematogaster modiglianii
Crematogaster adrepens
93 Crematogaster emeryana
Eurhopalothrix australis
Eurhopalothrix gravis
51 73 Talaridris mandibularis
Formicoxenus diversipilosus
Harpagoxenus sublaevis
100 Leptothorax muscorum complex
100 Liomyrmex gestroi
Metapone madagascarica
100 Metapone PG01
99 Myrmecina PG01
Myrmecina americana
100 Myrmecina graminicola
72 Nesomyrmex madecassus
Nesomyrmex MG01
100 Nesomyrmex MG08
Terataner bottegoi
Terataner MG02
Terataner MG27
71 Paratopula TH01
99 Romblonella scrobifera
Rotastruma recava
66 Rotastruma stenoceps
96 Dolopomyrmex pilatus
100 Monomorium latinode
74 Monomorium robustior
Monomorium criniceps
100 Monomorium destructor
90 Pogonomyrmex angustus
100 Hylomyrma blandiens
100 Pogonomyrmex striatinodus
Pogonomyrmex imberbiculus
Pogonomyrmex subdentatus
100 Pogonomyrmex vermiculatus
Manica bradleyi
100 Manica rubida
100 53 Myrmica kotokui
78 Myrmica rugiventris
Myrmica striolagaster
Myrmica tahoensis
100 Atta texana
Cyphomyrmex cornutus
63 Mycetarotes acutus
60 Cyphomyrmex longiscapus cf
72 Mycetosoritis hartmanni
Acromyrmex versicolor
Trachymyrmex arizonensis
98 Daceton armigerum
Lenomyrmex colwelli
63 Microdaceton tibialis
100 Colobostruma unicorna
Epopostruma monstrosa
Mesostruma browni
Orectognathus versicolor
98 Oxyopomyrmex santschii
54 100 Aphaenogaster swammerdami
Aphaenogaster pythia
52 Stenamma expolitum
70 Aphaenogaster araneoides
84 Aphaenogaster occidentalis
Messor denticornis
80 Messor wasmanni
Carebara nosindambo
73 Stegomyrmex manni
100 Diplomorium longipenne
100 Carebara vidua
100 75 Carebara urichi
100 &DUHEDUDDIILQLV
Carebara alperti
58 Carebara AU01
84 Solenopsis succinea
83 Solenopsis mameti
Solenopsis molesta
88 Solenopsis ZA02
85 Solenopsis fugax
100 Solenopsis xyloni
.HPSILGULVLQXVXDOLV
Tropidomyrmex elianae
Anisopheidole antipodum
Machomyrma dispar
99 Pheidole clydei
Pheidole fimbriata
Pheidole longispinosa
Pheidole pallidula
97 Pheidole rhea
Pheidole hyatti
Pheidole lucioccipitalis
57 Bariamyrma hispidula
Monomorium fisheri
96 Monomorium hanneli
Monomorium sechellense
100 Monomorium exiguum
92 Monomorium fieldi nr
Monomorium MG04
99 Monomorium pharaonis
Monomorium ergatogyna
Monomorium madecassum
7HWUDPRULXPYHQDWRU
Rhoptromyrmex transversinodis
97 Tetramorium caespitum cf
100 61 Tetramorium taylori
Tetramorium severini
Tetramorium spinosum
Strongylognathus testaceus
54 Tetramorium CF01
Tetramorium MG125
99 Tetramorium validiusculum
Anergates atratulus
Teleutomyrmex kutteri
6WUXPLJHQ\V ambatrix
Strumigenys exiguaevitae
6WUXPLJHQ\V olsoni
100 Strumigenys emmae
100 Strumigenys coveri
Strumigenys dicomas
87 Strumigenys rogeri
6WUXPLJHQ\V chiricahua
65 6WUXPLJHQ\V ocypete
72 6WUXPLJHQ\V membranifera
51 63 Strumigenys godeffroyi
6WUXPLJHQ\V nitens
Strumigenys chyzeri
54 98 6WUXPLJHQ\VPD[LOODULV
6WUXPLJHQ\V erynnes
100 6WUXPLJHQ\V ludovici
6WUXPLJHQ\V gundlachi
Strumigenys biolleyi
)LJXUH6$EG$

Apterogyna ZA01
Platythyrea mocquerysi
Paraponera clavata
77 81 Ankylomyrma coronacantha
100 Proceratium stictum
Tatuidris WDWXVLD
70 Tatuidris tatusiaB
Stigmatomma pallipes
Amblyopone australis
81 96 Tetraponera rufonigra
Acanthoponera minor
85 Rhytidoponera chalybaea
Aneuretus simoni
74 100 Dolichoderus pustulatus
77 Lasius californicus
98 Myrmelachista flavocotea
Myrmecia pyriformis
Nothomyrmecia macrops
100 Hylomyrma blandiens
65 Pogonomyrmex angustus
100 Pogonomyrmex striatinodus
87 Pogonomyrmex imberbiculus
Pogonomyrmex subdentatus
89 100 Pogonomyrmex vermiculatus
100 Manica bradleyi
90 Manica rubida
91 Myrmica kotokui
100 Myrmica rugiventris
Myrmica striolagaster
Myrmica tahoensis
Atopomyrmex mocquerysi
Austromorium flavigaster
Bariamyrma hispidula
Carebara urichi
Carebara vidua
Cyphoidris exalta
Dacatria templaris
Dacetinops ignotus cf
95 Dicroaspis KM01
Diplomorium longipenne
Dolopomyrmex pilatus
Kartidris sparsipila
Lophomyrmex ambiguus
Mayriella ebbei
Megalomyrmex silvestrii
Monomorium antarcticum
Monomorium ferodens
Monomorium latinode
Myrmisaraka brevis
Vitsika suspicax
&DUHEDUDDIILQLV
Podomyrma silvicola
Pristomyrmex orbiceps
Proatta butteli
Stegomyrmex manni
Tetheamyrma subspongia
66 Xenomyrmex floridanus
Adelomyrmex paratristani
57 Baracidris sitra
Adlerzia froggatti
100 Recurvidris TH01
Anillomyrma AFRC TZ 01
89 Anillomyrma decamera
Carebarella PA01
100 Solenopsis papuana
Dilobocondyla borneensis cf
100 Secostruma MY01
Melissotarsus insularis
91 100 Rhopalomastix rothneyi
Meranoplus castaneus
66 Meranoplus radamae
Monomorium brocha
95 Monomorium sechellense
Monomorium denticulatum
96 Oxyepoecus vezenyii
Monomorium hanneli
100 Tyrannomyrmex legatus
.HPSILGULVLQXVXDOLV
100 Tropidomyrmex elianae
Royidris notorthotenes
100 Royidris longiseta
Myrmicaria carinata
100 Myrmicaria exigua
MDODJLGULVMXJXP
61 MDODJLGULVEHOWL
Rogeria nevadensis
75 Rogeria stigmatica
91 Monomorium fisheri
Adelomyrmex SC01
100 Cryptomyrmex boltoni
87 Cardiocondyla MY01
Cardiocondyla mauritanica
82 Cardiocondyla thoracica
100 Ocymyrmex fortior cf
Cataulacus ebrardi
70 Cataulacus hispidulus
Eutetramorium mocquerysi
Huberia brounii
83 Perissomyrmex snyderi
99 Liomyrmex gestroi
Metapone madagascarica
100 Metapone PG01
100 Monomorium robustior
Monomorium criniceps
100 Monomorium destructor
65 Myrmecina PG01
Myrmecina americana
Myrmecina graminicola
99 Ancyridris polyrhachioides nr
100 Propodilobus pingorum
Lordomyrma bhutanensis cf
Lordomyrma desupra
55 Carebara nosindambo
100 Pristomyrmex punctatus
Carebara alperti
100 Carebara AU01
Nesomyrmex echinatinodis
77 Nesomyrmex wilda
100 Nesomyrmex madecassus
91 Nesomyrmex MG01
Nesomyrmex MG08
90 100 Acanthomyrmex ferox
96 Crematogaster acuta
99 Crematogaster sordidula
100 Crematogaster modiglianii
Crematogaster adrepens
100 Crematogaster emeryana
93 Monomorium fieldi nr
58 Monomorium ergatogyna
95 Monomorium madecassum
100 Monomorium pharaonis
Monomorium exiguum
Monomorium MG04
Solenopsis mameti
100 100 Solenopsis xyloni
Solenopsis fugax
77 Solenopsis ZA02
Solenopsis molesta
Solenopsis succinea
71 100 Paratopula TH01
73 Rotastruma recava
61 Rotastruma stenoceps
73 Poecilomyrma myrmecodiae
100 Romblonella scrobifera
92 Terataner bottegoi
Terataner MG02
Terataner MG27
100 Gauromyrmex acanthinus
59 100 Vombisidris bilongrudi
71 Harpagoxenus sublaevis
79 Formicoxenus diversipilosus
Leptothorax muscorum complex
97 Myrmoxenus bernardi
93 Temnothorax rugatulus
Chalepoxenus kutteri
Temnothorax poeyi
100 Stenamma striatulum
Aphaenogaster albisetosa
100 75 Messor andrei
Stenamma dyscheres
100 Stenamma expolitum
Aphaenogaster pythia
97 100 Aphaenogaster swammerdami
Goniomma blanci
99 Oxyopomyrmex santschii
100 Aphaenogaster araneoides
87 Aphaenogaster occidentalis
Messor denticornis
98 Messor wasmanni
97 Calyptomyrmex kaurus
Myrmicine genus26 PH02
Vollenhovia emeryi
86 Rhoptromyrmex transversinodis
96 Tetramorium MG125
Tetramorium severini
100 Tetramorium spinosum
100 Anergates atratulus
Strongylognathus testaceus
Teleutomyrmex kutteri
63 Tetramorium caespitum cf
99 Tetramorium taylori
100 Tetramorium CF01
7HWUDPRULXPYHQDWRU
Tetramorium validiusculum
Blepharidatta conops
Diaphoromyrma sofiae
Lachnomyrmex scrobiculatus
Octostruma EC01
100 Procryptocerus scabriusculus
Allomerus octoarticulatus
100 Wasmannia auropunctata
Cephalotes atratus
94 Cephalotes texanus
Phalacromyrmex fugax
64 Pilotrochus besmerus
86 Basiceros manni
Protalaridris armata
69 Rhopalothrix isthmica
100 Eurhopalothrix australis
Eurhopalothrix gravis
100 Talaridris mandibularis
94 Ochetomyrmex neopolitus
100 Ochetomyrmex semipolitus
Tranopelta subterranea
Tranopelta gilva
Pheidole fimbriata
69 Pheidole rhea
Pheidole hyatti
99 Pheidole longispinosa
75 Pheidole pallidula
63 Anisopheidole antipodum
76 Pheidole clydei
Machomyrma dispar
99 Pheidole lucioccipitalis
6WUXPLJHQ\V ambatrix
100 6WUXPLJHQ\V gundlachi
Strumigenys biolleyi
83 6WUXPLJHQ\V nitens
6WUXPLJHQ\V chiricahua
73 6WUXPLJHQ\V erynnes
Strumigenys exiguaevitae
6WUXPLJHQ\V ludovici
92 Strumigenys maxillaris
6WUXPLJHQ\V membranifera
6WUXPLJHQ\V ocypete
86 6WUXPLJHQ\V olsoni
Strumigenys chyzeri
Strumigenys emmae
100 Strumigenys godeffroyi
100 Strumigenys rogeri
Strumigenys coveri
Strumigenys dicomas
Acanthognathus ocellatus
Daceton armigerum
Kalathomyrmex emeryi
Lenomyrmex colwelli
Microdaceton tibialis
58 Myrmicocrypta infuscata cf
94 Orectognathus versicolor
Apterostigma auriculatum
73 Mycocepurus goeldii
98 Colobostruma unicorna
Epopostruma monstrosa
92 Mesostruma browni
Mycetarotes acutus
69 Mycetosoritis hartmanni
78 Cyphomyrmex cornutus
76 Cyphomyrmex longiscapus cf
100 Acromyrmex versicolor
Atta texana
Trachymyrmex arizonensis
)LJXUH6()D)

Apterogyna ZA01
Myrmecia pyriformis
100 Amblyopone australis
Platythyrea mocquerysi
77 Proceratium stictum
100 Stigmatomma pallipes
100 Ankylomyrma coronacantha
Tatuidris WDWXVLD
Tatuidris tatusiaB
Nothomyrmecia macrops
98 72 Paraponera clavata
Rhytidoponera chalybaea
75 Tetraponera rufonigra
74 Acanthoponera minor
100 Dolichoderus pustulatus
Lasius californicus
Myrmelachista flavocotea
Aneuretus simoni
Anillomyrma AFRC TZ 01
65 Anillomyrma decamera
Austromorium flavigaster
Bariamyrma hispidula
Carebarella PA01
Dolopomyrmex pilatus
Monomorium antarcticum
Monomorium brocha
Monomorium denticulatum
Monomorium hanneli
Monomorium latinode
Monomorium sechellense
Oxyepoecus vezenyii
80 Pogonomyrmex angustus
Rogeria nevadensis
55 Solenopsis papuana
Adelomyrmex paratristani
56 Tyrannomyrmex legatus
Megalomyrmex silvestrii
96 Rogeria stigmatica
.HPSILGULVLQXVXDOLV
100 Tropidomyrmex elianae
Myrmicaria carinata
100 Myrmicaria exigua
94 Cardiocondyla mauritanica
Cardiocondyla MY01
Cardiocondyla thoracica
61 Monomorium ferodens
99 Monomorium fisheri
72 Cryptomyrmex boltoni
Adelomyrmex SC01
77 Baracidris sitra
82 Hylomyrma blandiens
79 100 Pogonomyrmex striatinodus
54 Pogonomyrmex imberbiculus
Pogonomyrmex subdentatus
Pogonomyrmex vermiculatus
98 Monomorium exiguum
Monomorium fieldi nr
56 Monomorium MG04
Monomorium ergatogyna
Monomorium madecassum
100 Monomorium pharaonis
Solenopsis fugax
100 Solenopsis ZA02
100 Solenopsis mameti
99 Solenopsis molesta
Solenopsis succinea
Solenopsis xyloni
57 56 Acanthognathus ocellatus
Manica bradleyi
68 Manica rubida
100 Myrmica kotokui
Myrmica rugiventris
Myrmica striolagaster
Myrmica tahoensis
100 Calyptomyrmex kaurus
Cataulacus ebrardi
51 Cataulacus hispidulus
95 53 Nesomyrmex MG01
56 Nesomyrmex madecassus
90 Nesomyrmex MG08
Terataner bottegoi
Terataner MG02
Terataner MG27
Adlerzia froggatti
Aphaenogaster araneoides
Carebara vidua
Cyphoidris exalta
67 Dacatria templaris
Dacetinops ignotus cf
Diaphoromyrma sofiae
Dicroaspis KM01
Eutetramorium mocquerysi
Gauromyrmex acanthinus
Huberia brounii
Kartidris sparsipila
Mayriella ebbei
Melissotarsus insularis
MDODJLGULVMXJXP
MDODJLGULVEHOWL
Myrmisaraka brevis
Vitsika suspicax
Myrmicine genus26 PH02
Pheidole fimbriata
Proatta butteli
Procryptocerus scabriusculus
Recurvidris TH01
Rhopalomastix rothneyi
Stegomyrmex manni
Tetheamyrma subspongia
Vollenhovia emeryi
83 Vombisidris bilongrudi
Acanthomyrmex ferox
100 Diplomorium longipenne
Aphaenogaster albisetosa
65 Messor andrei
Aphaenogaster pythia
89 Aphaenogaster swammerdami
Carebara urichi
100 &DUHEDUDDIILQLV
Dilobocondyla borneensis cf
100 Secostruma MY01
Meranoplus castaneus
100 Meranoplus radamae
Royidris notorthotenes
100 Royidris longiseta
Nesomyrmex echinatinodis
77 Nesomyrmex wilda
Perissomyrmex snyderi
99 Podomyrma silvicola
Pristomyrmex orbiceps
97 Pristomyrmex punctatus
57 Carebara nosindambo
Carebara alperti
100 Carebara AU01
Formicoxenus diversipilosus
Harpagoxenus sublaevis
72 Leptothorax muscorum complex
85 Liomyrmex gestroi
Metapone madagascarica
97 Metapone PG01
Monomorium criniceps
Monomorium destructor
98 Monomorium robustior
98 Stenamma expolitum
Stenamma dyscheres
66 Stenamma striatulum
60 Allomerus octoarticulatus
60 Wasmannia auropunctata
Blepharidatta conops
Lachnomyrmex scrobiculatus
96 Chalepoxenus kutteri
Myrmoxenus bernardi
Temnothorax poeyi
Temnothorax rugatulus
70 Poecilomyrma myrmecodiae
100 Romblonella scrobifera
Rotastruma recava
Rotastruma stenoceps
57 54 Lophomyrmex ambiguus
99 Ancyridris polyrhachioides nr
91 Propodilobus pingorum
Lordomyrma bhutanensis cf
100 Lordomyrma desupra
Goniomma blanci
99 Oxyopomyrmex santschii
87 100 Aphaenogaster occidentalis
Messor denticornis
58 Messor wasmanni
Ocymyrmex fortior cf
51 Paratopula TH01
100 97 Myrmecina PG01
Myrmecina americana
Myrmecina graminicola
82 100 Atopomyrmex mocquerysi
Crematogaster acuta
77 Crematogaster sordidula
99 Crematogaster modiglianii
Crematogaster adrepens
100 Crematogaster emeryana
Cephalotes atratus
80 100 Cephalotes texanus
100 Ochetomyrmex neopolitus
100 Ochetomyrmex semipolitus
Tranopelta subterranea
Tranopelta gilva
Basiceros manni
98 Octostruma EC01
Protalaridris armata
95 Rhopalothrix isthmica
97 Eurhopalothrix australis
Eurhopalothrix gravis
Talaridris mandibularis
82 Daceton armigerum
65 Lenomyrmex colwelli
Microdaceton tibialis
100 Orectognathus versicolor
94 Colobostruma unicorna
Epopostruma monstrosa
Mesostruma browni
96 Pheidole clydei
Pheidole hyatti
67 Pheidole rhea
Anisopheidole antipodum
59 Machomyrma dispar
Pheidole longispinosa
Pheidole lucioccipitalis
Pheidole pallidula
82 Xenomyrmex floridanus
Apterostigma auriculatum
64 Myrmicocrypta infuscata cf
54 Mycocepurus goeldii
96 Kalathomyrmex emeryi
51 Mycetarotes acutus
Mycetosoritis hartmanni
99 Cyphomyrmex longiscapus cf
51 100 Cyphomyrmex cornutus
61 Trachymyrmex arizonensis
Acromyrmex versicolor
Atta texana
7HWUDPRULXPYHQDWRU
Rhoptromyrmex transversinodis
100 Tetramorium validiusculum
Tetramorium CF01
73 Tetramorium taylori
73 73 Tetramorium MG125
Tetramorium severini
97 Tetramorium spinosum
67 Anergates atratulus
100 Teleutomyrmex kutteri
Strongylognathus testaceus
98 Tetramorium caespitum cf
Phalacromyrmex fugax
Pilotrochus besmerus
6WUXPLJHQ\V ambatrix
99 95 Strumigenys rogeri
6WUXPLJHQ\V erynnes
69 6WUXPLJHQ\V ludovici
Strumigenys coveri
93 82 Strumigenys dicomas
6WUXPLJHQ\V gundlachi
6WUXPLJHQ\V nitens
Strumigenys biolleyi
Strumigenys exiguaevitae
6WUXPLJHQ\V olsoni
78 82 Strumigenys godeffroyi
6WUXPLJHQ\V chiricahua
75 6WUXPLJHQ\V membranifera
61 6WUXPLJHQ\V ocypete
Strumigenys maxillaris
Strumigenys chyzeri
Strumigenys emmae
)LJXUH6/:5K

Apterogyna ZA01
Acanthognathus ocellatus
Acanthomyrmex ferox
Adlerzia froggatti
Atopomyrmex mocquerysi
Calyptomyrmex kaurus
Cyphoidris exalta
Dacetinops ignotus cf
Daceton armigerum
Diaphoromyrma sofiae
Dicroaspis KM01
Diplomorium longipenne
Kartidris sparsipila
Lachnomyrmex scrobiculatus
Lenomyrmex colwelli
Microdaceton tibialis
Orectognathus versicolor
Perissomyrmex snyderi
Recurvidris TH01
Rogeria stigmatica
Stegomyrmex manni
96 Vombisidris bilongrudi
Dacatria templaris
99 Proatta butteli
Huberia brounii
100 Podomyrma silvicola
Meranoplus castaneus
83 Meranoplus radamae
Myrmicine genus26 PH02
100 Vollenhovia emeryi
Pristomyrmex orbiceps
100 Pristomyrmex punctatus
100 Blepharidatta conops
Allomerus octoarticulatus
87 Wasmannia auropunctata
99 Apterostigma auriculatum
Mycocepurus goeldii
98 Myrmicocrypta infuscata cf
100 Colobostruma unicorna
Epopostruma monstrosa
68 Mesostruma browni
Gauromyrmex acanthinus
Mayriella ebbei
72 Tetheamyrma subspongia
99 Lophomyrmex ambiguus
Melissotarsus insularis
100 Rhopalomastix rothneyi
100 Myrmecina PG01
Myrmecina americana
100 Myrmecina graminicola
54 Terataner MG02
Terataner bottegoi
100 Terataner MG27
77 Lordomyrma bhutanensis cf
51 Lordomyrma desupra
Ancyridris polyrhachioides nr
100 Propodilobus pingorum
56 Temnothorax poeyi
91 Temnothorax rugatulus
Chalepoxenus kutteri
100 Myrmoxenus bernardi
100 Ochetomyrmex neopolitus
98 Ochetomyrmex semipolitus
Tranopelta subterranea
100 Tranopelta gilva
Cephalotes atratus
90 Cephalotes texanus
97 100 Procryptocerus scabriusculus
Dilobocondyla borneensis cf
Secostruma MY01
97 60 Eutetramorium mocquerysi
100 Malagidris jugum
100 MDODJLGULVEHOWL
Myrmisaraka brevis
Vitsika suspicax
78 92 Paratopula TH01
56 Poecilomyrma myrmecodiae
100 Romblonella scrobifera
Rotastruma recava
Rotastruma stenoceps
99 Carebara vidua
100 100 Carebara nosindambo
53 Carebara alperti
100 Carebara AU01
Carebara urichi
&DUHEDUDDIILQLV
Crematogaster modiglianii
57 96 Ocymyrmex fortior cf
Crematogaster acuta
71 Crematogaster sordidula
Crematogaster adrepens
95 Crematogaster emeryana
78 Kalathomyrmex emeryi
90 Mycetarotes acutus
100 Mycetosoritis hartmanni
96 Cyphomyrmex cornutus
100 Cyphomyrmex longiscapus cf
92 Trachymyrmex arizonensis
Acromyrmex versicolor
Atta texana
100 Pheidole rhea
57 Pheidole fimbriata
Pheidole longispinosa
100 53 Pheidole lucioccipitalis
Anisopheidole antipodum
70 Machomyrma dispar
94 Pheidole pallidula
Pheidole clydei
Pheidole hyatti
Basiceros manni
Octostruma EC01
58 56 Protalaridris armata
100 Rhopalothrix isthmica
Cataulacus ebrardi
91 Cataulacus hispidulus
99 Eurhopalothrix australis
Eurhopalothrix gravis
99 Talaridris mandibularis
100 Xenomyrmex floridanus
Formicoxenus diversipilosus
94 Harpagoxenus sublaevis
100 Leptothorax muscorum complex
Nesomyrmex echinatinodis
80 Nesomyrmex wilda
100 88 Nesomyrmex MG08
Nesomyrmex madecassus
68 Nesomyrmex MG01
100 Liomyrmex gestroi
Metapone madagascarica
70 100 Metapone PG01
75 Monomorium destructor
Monomorium criniceps
54 100 Monomorium robustior
61 Royidris notorthotenes
100 Royidris longiseta
100 Cardiocondyla mauritanica
Cardiocondyla MY01
Cardiocondyla thoracica
93 Tetramorium spinosum
7HWUDPRULXPYHQDWRU
100 60 Tetramorium validiusculum
Rhoptromyrmex transversinodis
100 Tetramorium MG125
Tetramorium CF01
70 Tetramorium taylori
100 Tetramorium severini
94 Tetramorium caespitum cf
99 Strongylognathus testaceus
Anergates atratulus
100 Teleutomyrmex kutteri
Aphaenogaster albisetosa
99 Messor andrei
100 59 Stenamma expolitum
100 Stenamma dyscheres
72 Stenamma striatulum
100 Aphaenogaster araneoides
Aphaenogaster pythia
97 100 Aphaenogaster swammerdami
100 Goniomma blanci
100 Oxyopomyrmex santschii
100 Aphaenogaster occidentalis
Messor denticornis
99 Messor wasmanni
Phalacromyrmex fugax
61 Pilotrochus besmerus
6WUXPLJHQ\V ambatrix
6WUXPLJHQ\V erynnes
97 6WUXPLJHQ\V ludovici
92 Strumigenys biolleyi
6WUXPLJHQ\V gundlachi
80 73 6WUXPLJHQ\V nitens
Strumigenys exiguaevitae
100 Strumigenys maxillaris
Strumigenys coveri
Strumigenys dicomas
Strumigenys rogeri
56 6WUXPLJHQ\V olsoni
6WUXPLJHQ\V chiricahua
100 6WUXPLJHQ\V membranifera
6WUXPLJHQ\V ocypete
66 Strumigenys godeffroyi
Strumigenys chyzeri
Strumigenys emmae
99 Hylomyrma blandiens
Pogonomyrmex angustus
93 100 Pogonomyrmex striatinodus
93 Pogonomyrmex imberbiculus
Pogonomyrmex subdentatus
56 100 Pogonomyrmex vermiculatus
Manica bradleyi
100 Manica rubida
100 98 Myrmica kotokui
84 Myrmica rugiventris
Myrmica striolagaster
63 Myrmica tahoensis
100 Aneuretus simoni
Acanthoponera minor
98 Rhytidoponera chalybaea
Lasius californicus
64 71 Myrmelachista flavocotea
92 Dolichoderus pustulatus
100 Tetraponera rufonigra
Myrmecia pyriformis
Nothomyrmecia macrops
61 Platythyrea mocquerysi
57 56 Stigmatomma pallipes
Amblyopone australis
70 Proceratium stictum
100 Paraponera clavata
100 Ankylomyrma coronacantha
Tatuidris WDWXVLD
Tatuidris tatusiaB
Austromorium flavigaster
Bariamyrma hispidula
Monomorium sechellense
61 Oxyepoecus vezenyii
Megalomyrmex silvestrii
70 Monomorium denticulatum
57 Monomorium antarcticum
Monomorium fisheri
87 Rogeria nevadensis
100 Adelomyrmex paratristani
98 Baracidris sitra
Adelomyrmex SC01
90 100 Cryptomyrmex boltoni
Carebarella PA01
100 99 Solenopsis papuana
100 Solenopsis fugax
93 Solenopsis ZA02
99 Solenopsis xyloni
100 Solenopsis mameti
Solenopsis molesta
Solenopsis succinea
Monomorium ferodens
86 Monomorium hanneli
58 Monomorium brocha
Monomorium latinode
51 59 100 Tyrannomyrmex legatus
89 Anillomyrma AFRC TZ 01
100 Anillomyrma decamera
Myrmicaria carinata
100 Myrmicaria exigua
.HPSILGULVLQXVXDOLV
53 Tropidomyrmex elianae
99 Dolopomyrmex pilatus
Monomorium ergatogyna
100 Monomorium madecassum
63 Monomorium MG04
58 Monomorium fieldi nr
Monomorium exiguum
Monomorium pharaonis
)LJXUH6$UJ.

Apterogyna ZA01
100 100 Platythyrea mocquerysi
Amblyopone australis
93 Stigmatomma pallipes
57 Paraponera clavata
100 Proceratium stictum
100 Ankylomyrma coronacantha
Tatuidris WDWXVLD
92 Tatuidris tatusiaB
Acanthoponera minor
100 Rhytidoponera chalybaea
Lasius californicus
Myrmelachista flavocotea
84 Aneuretus simoni
85 Dolichoderus pustulatus
100 Tetraponera rufonigra
Myrmecia pyriformis
100 Nothomyrmecia macrops
Manica bradleyi
100 Manica rubida
100 Myrmica kotokui
100 Myrmica rugiventris
Myrmica striolagaster
Myrmica tahoensis
100 Hylomyrma blandiens
86 66 Pogonomyrmex angustus
Pogonomyrmex striatinodus
100 55 Pogonomyrmex imberbiculus
Pogonomyrmex subdentatus
Pogonomyrmex vermiculatus
100 Stenamma dyscheres
Aphaenogaster albisetosa
100 70 Messor andrei
Stenamma expolitum
100 Stenamma striatulum
Aphaenogaster pythia
99 Aphaenogaster swammerdami
85 100 Aphaenogaster araneoides
98 Goniomma blanci
100 Oxyopomyrmex santschii
100 Aphaenogaster occidentalis
Messor denticornis
98 Messor wasmanni
Acanthomyrmex ferox
Adlerzia froggatti
Dacatria templaris
Dacetinops ignotus cf
Eutetramorium mocquerysi
Huberia brounii
Kartidris sparsipila
Mayriella ebbei
Ocymyrmex fortior cf
Podomyrma silvicola
Proatta butteli
Rotastruma recava
Tetheamyrma subspongia
100 Xenomyrmex floridanus
Melissotarsus insularis
100 Rhopalomastix rothneyi
Meranoplus castaneus
100 Meranoplus radamae
Royidris notorthotenes
78 Royidris longiseta
Myrmicine genus26 PH02
76 Vollenhovia emeryi
90 100 Liomyrmex gestroi
Metapone madagascarica
100 Metapone PG01
71 Monomorium robustior
Monomorium criniceps
100 Monomorium destructor
81 Myrmecina PG01
Myrmecina americana
68 Myrmecina graminicola
100 Perissomyrmex snyderi
Pristomyrmex orbiceps
100 Pristomyrmex punctatus
99 Terataner bottegoi
Terataner MG02
Terataner MG27
99 Lordomyrma desupra
79 Propodilobus pingorum
Ancyridris polyrhachioides nr
100 Lordomyrma bhutanensis cf
Malagidris jugum
89
96 94 MDODJLGULVEHOWL
Myrmisaraka brevis
100 Vitsika suspicax
100 Crematogaster acuta
100 Crematogaster sordidula
100 Crematogaster modiglianii
Crematogaster adrepens
100 Crematogaster emeryana
Nesomyrmex echinatinodis
100 Nesomyrmex wilda
100 79 Nesomyrmex madecassus
Nesomyrmex MG01
Nesomyrmex MG08
71 Recurvidris TH01
92 Diplomorium longipenne
99 100 Carebara vidua
84 Carebara urichi
65 &DUHEDUDDIILQLV
100 Carebara nosindambo
Carebara alperti
100 Carebara AU01
Cataulacus ebrardi
98 Cataulacus hispidulus
98 94 100 Lophomyrmex ambiguus
Dilobocondyla borneensis cf
98 Secostruma MY01
100 Paratopula TH01
Poecilomyrma myrmecodiae
Romblonella scrobifera
73 Rotastruma stenoceps
100 Atopomyrmex mocquerysi
72 93 Harpagoxenus sublaevis
Formicoxenus diversipilosus
88 Leptothorax muscorum complex
88 Gauromyrmex acanthinus
100 Vombisidris bilongrudi
95 Chalepoxenus kutteri
90 Myrmoxenus bernardi
Temnothorax poeyi
Temnothorax rugatulus
Calyptomyrmex kaurus
Cyphoidris exalta
100 Dicroaspis KM01
58 93 Cardiocondyla mauritanica
Cardiocondyla MY01
100 Cardiocondyla thoracica
Tetramorium CF01
91 Tetramorium taylori
100 89 Tetramorium caespitum cf
100 93 Strongylognathus testaceus
Anergates atratulus
86 Teleutomyrmex kutteri
7HWUDPRULXPYHQDWRU
60 Tetramorium validiusculum
57 Tetramorium severini
89 Tetramorium spinosum
Rhoptromyrmex transversinodis
Tetramorium MG125
Bariamyrma hispidula
Dolopomyrmex pilatus
Megalomyrmex silvestrii
Monomorium antarcticum
Monomorium denticulatum
Monomorium sechellense
Oxyepoecus vezenyii
Rogeria nevadensis
Rogeria stigmatica
Stegomyrmex manni
100 Tyrannomyrmex legatus
Anillomyrma AFRC TZ 01
93 Anillomyrma decamera
Austromorium flavigaster
69 Monomorium brocha
Monomorium ferodens
99 51 Monomorium fisheri
Monomorium hanneli
100 Monomorium latinode
Kempfidris inusualis
100 Tropidomyrmex elianae
Myrmicaria carinata
73 Myrmicaria exigua
53 Adelomyrmex paratristani
51 Baracidris sitra
Adelomyrmex SC01
Cryptomyrmex boltoni
100 Monomorium ergatogyna
Monomorium pharaonis
58 Monomorium fieldi nr
100 Monomorium madecassum
Monomorium exiguum
100 Monomorium MG04
Carebarella PA01
100 100 Solenopsis papuana
100 Solenopsis fugax
87 Solenopsis ZA02
100 Solenopsis mameti
83 Solenopsis xyloni
Solenopsis molesta
57 Solenopsis succinea
Acanthognathus ocellatus
Blepharidatta conops
Diaphoromyrma sofiae
Lachnomyrmex scrobiculatus
100 Lenomyrmex colwelli
Allomerus octoarticulatus
100 Wasmannia auropunctata
76 Apterostigma auriculatum
Mycocepurus goeldii
100 Myrmicocrypta infuscata cf
100 Procryptocerus scabriusculus
Cephalotes atratus
100 Cephalotes texanus
100 Orectognathus versicolor
100 Colobostruma unicorna
Epopostruma monstrosa
100 Mesostruma browni
79 Ochetomyrmex neopolitus
100 Ochetomyrmex semipolitus
Tranopelta subterranea
Tranopelta gilva
100 100 Eurhopalothrix australis
88 Eurhopalothrix gravis
89 Talaridris mandibularis
99 96 Basiceros manni
95 Octostruma EC01
Protalaridris armata
65 Rhopalothrix isthmica
Daceton armigerum
68 Kalathomyrmex emeryi
100 77 Mycetosoritis hartmanni
Cyphomyrmex cornutus
66 Mycetarotes acutus
100 Cyphomyrmex longiscapus cf
Acromyrmex versicolor
Atta texana
Trachymyrmex arizonensis
61 Microdaceton tibialis
99 Pheidole rhea
61 Pheidole fimbriata
100 Anisopheidole antipodum
Pheidole longispinosa
87 Machomyrma dispar
97 Pheidole lucioccipitalis
56 Pheidole pallidula
Pheidole clydei
100 Pheidole hyatti
Phalacromyrmex fugax
100 Pilotrochus besmerus
56 58 6WUXPLJHQ\V gundlachi
6WUXPLJHQ\V nitens
100 Strumigenys biolleyi
100 6WUXPLJHQ\V ambatrix
6WUXPLJHQ\V erynnes
55 79 6WUXPLJHQ\V ludovici
88 Strumigenys rogeri
Strumigenys coveri
93 Strumigenys dicomas
Strumigenys maxillaris
100 6WUXPLJHQ\V olsoni
99 Strumigenys exiguaevitae
75 91 6WUXPLJHQ\V membranifera
6WUXPLJHQ\V chiricahua
87 6WUXPLJHQ\V ocypete
100 Strumigenys emmae
Strumigenys chyzeri
Strumigenys godeffroyi
)LJXUH67RS

85 Apterogyna ZA01
100 Acanthoponera minor
Myrmecia pyriformis
78 Nothomyrmecia macrops
94 Tetraponera rufonigra
Aneuretus simoni
Dolichoderus pustulatus
100 83 Proceratium stictum
Amblyopone australis
76 Stigmatomma pallipes
95 Platythyrea mocquerysi
66 100 Paraponera clavata
100 Ankylomyrma coronacantha
Tatuidris WDWXVLD
93 100 Tatuidris tatusiaB
Lasius californicus
100 Myrmelachista flavocotea
Manica bradleyi
100 Manica rubida
100 Myrmica kotokui
84 100 Myrmica rugiventris
Myrmica striolagaster
100 Myrmica tahoensis
97 Pogonomyrmex angustus
100 Hylomyrma blandiens
97 Pogonomyrmex striatinodus
86 54 Pogonomyrmex imberbiculus
Pogonomyrmex subdentatus
Pogonomyrmex vermiculatus
Rhytidoponera chalybaea
100 Aphaenogaster araneoides
Aphaenogaster albisetosa
100 99 Messor andrei
63 100 Stenamma expolitum
54 92 Stenamma dyscheres
100 Stenamma striatulum
Aphaenogaster pythia
68 100 Aphaenogaster swammerdami
100 Goniomma blanci
100 Oxyopomyrmex santschii
100 Aphaenogaster occidentalis
Messor denticornis
Messor wasmanni
Microdaceton tibialis
100 Myrmicine genus26 PH02
Protalaridris armata
61 Rhopalothrix isthmica
67 74 Blepharidatta conops
90 100 Tetheamyrma subspongia
Allomerus octoarticulatus
Wasmannia auropunctata
51 Vollenhovia emeryi
100 Calyptomyrmex kaurus
93 Tetramorium CF01
Tetramorium taylori
100 100 Rhoptromyrmex transversinodis
100 Tetramorium MG125
Tetramorium severini
100 Tetramorium spinosum
7HWUDPRULXPYHQDWRU
88 Tetramorium validiusculum
100 80 Tetramorium caespitum cf
99 Strongylognathus testaceus
Anergates atratulus
93 96 Teleutomyrmex kutteri
Dolopomyrmex pilatus
92 Rogeria stigmatica
Bariamyrma hispidula
100 100 Rogeria nevadensis
85 Austromorium flavigaster
99 Monomorium antarcticum
Monomorium denticulatum
67 50 100 Oxyepoecus vezenyii
Kempfidris inusualis
71 100 Tropidomyrmex elianae
Carebarella PA01
100 100 Solenopsis papuana
100 Solenopsis fugax
100 Solenopsis ZA02
100 100 Solenopsis mameti
81 Solenopsis xyloni
Solenopsis molesta
Solenopsis succinea
Megalomyrmex silvestrii
92 99 Baracidris sitra
Adelomyrmex SC01
71 85 Cryptomyrmex boltoni
99 Adelomyrmex paratristani
100 100 Monomorium madecassum
97 Monomorium exiguum
100 51 Monomorium MG04
56 Monomorium ergatogyna
Monomorium fieldi nr
Monomorium pharaonis
66 Monomorium ferodens
86 Monomorium fisheri
Monomorium hanneli
94 100 Monomorium sechellense
93 Myrmicaria carinata
74 Myrmicaria exigua
96 Monomorium brocha
Monomorium latinode
100 Tyrannomyrmex legatus
Anillomyrma AFRC TZ 01
100 Anillomyrma decamera
100 Ochetomyrmex neopolitus
100 Ochetomyrmex semipolitus
Tranopelta subterranea
61 100 Tranopelta gilva
97 Octostruma EC01
99 Basiceros manni
100 100 Eurhopalothrix australis
Eurhopalothrix gravis
100 Talaridris mandibularis
Phalacromyrmex fugax
100 Pilotrochus besmerus
98 6WUXPLJHQ\V ambatrix
100 100 6WUXPLJHQ\V nitens
6WUXPLJHQ\V gundlachi
98 100 Strumigenys biolleyi
100 Strumigenys rogeri
Strumigenys coveri
100 100 Strumigenys dicomas
6WUXPLJHQ\V erynnes
82 6WUXPLJHQ\V ludovici
100 Strumigenys maxillaris
6WUXPLJHQ\V chiricahua
52 6WUXPLJHQ\V membranifera
61 6WUXPLJHQ\V ocypete
Strumigenys exiguaevitae
78 6WUXPLJHQ\V olsoni
93 Strumigenys godeffroyi
Strumigenys chyzeri
Strumigenys emmae
Lachnomyrmex scrobiculatus
97 Stegomyrmex manni
100 Procryptocerus scabriusculus
Cephalotes atratus
55 Cephalotes texanus
100 Pheidole fimbriata
100 Pheidole rhea
100 Pheidole hyatti
Anisopheidole antipodum
76 Machomyrma dispar
77 Pheidole clydei
88 Pheidole longispinosa
Pheidole lucioccipitalis
Pheidole pallidula
Acanthognathus ocellatus
Daceton armigerum
98 Diaphoromyrma sofiae
100 Lenomyrmex colwelli
100 Orectognathus versicolor
52 100 Colobostruma unicorna
Epopostruma monstrosa
63 100 Mesostruma browni
Mycocepurus goeldii
100 Myrmicocrypta infuscata cf
96 Apterostigma auriculatum
99 Kalathomyrmex emeryi
100 Mycetarotes acutus
72 96 Mycetosoritis hartmanni
100 Cyphomyrmex cornutus
100 Cyphomyrmex longiscapus cf
61 Atta texana
Acromyrmex versicolor
Trachymyrmex arizonensis
Adlerzia froggatti
Dacetinops ignotus cf
Mayriella ebbei
52 Perissomyrmex snyderi
Cyphoidris exalta
95 Dicroaspis KM01
Dacatria templaris
100 Proatta butteli
Dilobocondyla borneensis cf
58 Secostruma MY01
Kartidris sparsipila
100 Lophomyrmex ambiguus
Melissotarsus insularis
100 Rhopalomastix rothneyi
100 Monomorium robustior
Monomorium criniceps
100 Monomorium destructor
77 Ancyridris polyrhachioides nr
55 Lordomyrma desupra
Lordomyrma bhutanensis cf
100 Propodilobus pingorum
64 Meranoplus castaneus
100 Meranoplus radamae
Pristomyrmex orbiceps
Pristomyrmex punctatus
98 Paratopula TH01
Poecilomyrma myrmecodiae
100 Romblonella scrobifera
Rotastruma recava
Rotastruma stenoceps
96 Diplomorium longipenne
98 Carebara vidua
74 100 Carebara nosindambo
91 Carebara alperti
100 Carebara AU01
Carebara urichi
77 100 &DUHEDUDDIILQLV
100 Myrmecina PG01
Myrmecina americana
57 97 Myrmecina graminicola
100 Crematogaster acuta
100 Crematogaster sordidula
100 Crematogaster modiglianii
Crematogaster adrepens
Crematogaster emeryana
99 100 Atopomyrmex mocquerysi
Terataner bottegoi
89 Terataner MG02
100 Terataner MG27
Cataulacus ebrardi
70 Cataulacus hispidulus
100 100 Ocymyrmex fortior cf
99 Cardiocondyla mauritanica
Cardiocondyla MY01
Cardiocondyla thoracica
63 Acanthomyrmex ferox
Podomyrma silvicola
58 Xenomyrmex floridanus
100 Huberia brounii
53 100 Liomyrmex gestroi
Metapone madagascarica
Metapone PG01
100 100 Eutetramorium mocquerysi
Royidris notorthotenes
100 Royidris longiseta
100 Malagidris jugum
100 MDODJLGULVEHOWL
Myrmisaraka brevis
Vitsika suspicax
54 Recurvidris TH01
74 100 Vombisidris bilongrudi
100 Nesomyrmex echinatinodis
100 Nesomyrmex wilda
79 55 Nesomyrmex madecassus
Nesomyrmex MG01
Nesomyrmex MG08
100 Gauromyrmex acanthinus
79 89 Harpagoxenus sublaevis
Formicoxenus diversipilosus
69 Leptothorax muscorum complex
100 Chalepoxenus kutteri
65 Myrmoxenus bernardi
Temnothorax poeyi
Temnothorax rugatulus
)LJXUH6UB[

100 Apterogyna ZA01


Myrmicine genus26 PH02
Vollenhovia emeryi
Calyptomyrmex kaurus
100 Melissotarsus insularis
96 Cardiocondyla mauritanica
Cardiocondyla MY01
88 Cardiocondyla thoracica
Tetramorium CF01
89 Tetramorium taylori
100 Rhoptromyrmex transversinodis
99 Tetramorium MG125
84 100 Anergates atratulus
94 Teleutomyrmex kutteri
Strongylognathus testaceus
96 Tetramorium caespitum cf
7HWUDPRULXPYHQDWRU
86
88 100 Tetramorium validiusculum
Tetramorium severini
Tetramorium spinosum
Acanthomyrmex ferox
Adlerzia froggatti
Atopomyrmex mocquerysi
Dacatria templaris
Dacetinops ignotus cf
Dicroaspis KM01
Kartidris sparsipila
Mayriella ebbei
Paratopula TH01
Perissomyrmex snyderi
Proatta butteli
Recurvidris TH01
Rhopalomastix rothneyi
85 Tetheamyrma subspongia
100 Xenomyrmex floridanus
Cataulacus ebrardi
100 Cataulacus hispidulus
Dilobocondyla borneensis cf
100 Secostruma MY01
Pristomyrmex orbiceps
100 Pristomyrmex punctatus
77 Monomorium criniceps
Monomorium destructor
100 Monomorium robustior
98 Terataner bottegoi
Terataner MG02
100 Terataner MG27
66 Ancyridris polyrhachioides nr
Lordomyrma bhutanensis cf
Lordomyrma desupra
71 Propodilobus pingorum
100 Cyphoidris exalta
96 Myrmecina PG01
Myrmecina americana
Myrmecina graminicola
58 Lophomyrmex ambiguus
100 Ocymyrmex fortior cf
Meranoplus castaneus
Meranoplus radamae
100 Poecilomyrma myrmecodiae
83 100 Romblonella scrobifera
Rotastruma recava
96 Rotastruma stenoceps
Crematogaster acuta
98 Crematogaster sordidula
79 71 Crematogaster modiglianii
Crematogaster adrepens
Crematogaster emeryana
97 Huberia brounii
100 Podomyrma silvicola
100 Liomyrmex gestroi
Metapone madagascarica
100 Metapone PG01
Nesomyrmex echinatinodis
91 Nesomyrmex wilda
100 76 Nesomyrmex MG01
Nesomyrmex madecassus
Nesomyrmex MG08
97 Diplomorium longipenne
100 Carebara vidua
57 100 Carebara nosindambo
55 Carebara alperti
100 Carebara AU01
Carebara urichi
&DUHEDUDDIILQLV
61 Eutetramorium mocquerysi
Malagidris jugum
69 MyDODJLGULVEHOWL
Myrmisaraka brevis
100 Vitsika suspicax
Royidris notorthotenes
59 Royidris longiseta
99 Vombisidris bilongrudi
97 Formicoxenus diversipilosus
Harpagoxenus sublaevis
90 Leptothorax muscorum complex
100 Gauromyrmex acanthinus
78 Myrmoxenus bernardi
Chalepoxenus kutteri
Temnothorax poeyi
100 Temnothorax rugatulus
Stenamma dyscheres
Stenamma expolitum
66 100 Stenamma striatulum
Aphaenogaster albisetosa
65 100 Messor andrei
Aphaenogaster pythia
95 Aphaenogaster swammerdami
64 100 Aphaenogaster araneoides
Goniomma blanci
71 Oxyopomyrmex santschii
100 Aphaenogaster occidentalis
Messor denticornis
87 Messor wasmanni
100 Pogonomyrmex angustus
68 Hylomyrma blandiens
87 Pogonomyrmex imberbiculus
Pogonomyrmex striatinodus
Pogonomyrmex subdentatus
100 Pogonomyrmex vermiculatus
Manica bradleyi
90 100 Manica rubida
62 63 Myrmica kotokui
72 Myrmica rugiventris
Myrmica striolagaster
99 Myrmica tahoensis
Acanthoponera minor
90 100 Rhytidoponera chalybaea
Lasius californicus
58 81 Myrmelachista flavocotea
100 Tetraponera rufonigra
79 Aneuretus simoni
Dolichoderus pustulatus
78 Myrmecia pyriformis
53 Nothomyrmecia macrops
91 100 Platythyrea mocquerysi
100 Ankylomyrma coronacantha
88 Tatuidris WDWXVLD
Tatuidris tatusiaB
89 Amblyopone australis
97 Proceratium stictum
Paraponera clavata
Stigmatomma pallipes
Dolopomyrmex pilatus
Rogeria nevadensis
Stegomyrmex manni
Adelomyrmex paratristani
Austromorium flavigaster
Baracidris sitra
Bariamyrma hispidula
99 Megalomyrmex silvestrii
Monomorium antarcticum
Monomorium brocha
Monomorium denticulatum
Monomorium hanneli
Monomorium latinode
Monomorium sechellense
Oxyepoecus vezenyii
Rogeria stigmatica
66 91 Tyrannomyrmex legatus
Adelomyrmex SC01
100 Cryptomyrmex boltoni
Anillomyrma AFRC TZ 01
85 Anillomyrma decamera
Monomorium ferodens
100 Monomorium fisheri
Myrmicaria carinata
Myrmicaria exigua
100 Monomorium pharaonis
80 61 Monomorium exiguum
Monomorium ergatogyna
75 Monomorium madecassum
Monomorium fieldi nr
100 Monomorium MG04
Kempfidris inusualis
99 Tropidomyrmex elianae
90 Carebarella PA01
72 Solenopsis papuana
Solenopsis fugax
100 Solenopsis ZA02
94 100 Solenopsis mameti
100 Solenopsis xyloni
Solenopsis molesta
Solenopsis succinea
Acanthognathus ocellatus
Apterostigma auriculatum
Daceton armigerum
60 Lachnomyrmex scrobiculatus
Diaphoromyrma sofiae
71 Lenomyrmex colwelli
Mycocepurus goeldii
85 Myrmicocrypta infuscata cf
100 Blepharidatta conops
Allomerus octoarticulatus
100 Wasmannia auropunctata
100 Procryptocerus scabriusculus
Cephalotes atratus
100 Cephalotes texanus
97 Ochetomyrmex neopolitus
100 Ochetomyrmex semipolitus
Tranopelta subterranea
Tranopelta gilva
88 Microdaceton tibialis
97 Colobostruma unicorna
100 Orectognathus versicolor
Epopostruma monstrosa
54 Mesostruma browni
Cyphomyrmex longiscapus cf
78 Kalathomyrmex emeryi
Mycetarotes acutus
58 Mycetosoritis hartmanni
100 Cyphomyrmex cornutus
61 Trachymyrmex arizonensis
Acromyrmex versicolor
Atta texana
Pheidole fimbriata
100 Pheidole rhea
Machomyrma dispar
84 Pheidole clydei
70 Pheidole longispinosa
Anisopheidole antipodum
99 Pheidole hyatti
Pheidole lucioccipitalis
72 Pheidole pallidula
100 91 Basiceros manni
Protalaridris armata
81 Rhopalothrix isthmica
100 Octostruma EC01
100 Eurhopalothrix australis
Eurhopalothrix gravis
80 89 Talaridris mandibularis
Phalacromyrmex fugax
Pilotrochus besmerus
100 83 6WUXPLJHQ\V ambatrix
83 6WUXPLJHQ\V nitens
Strumigenys gundlachi
Strumigenys biolleyi
100 Strumigenys chiricahua
Strumigenys exiguaevitae
Strumigenys maxillaris
Strumigenys membranifera
51 Strumigenys ocypete
98 Strumigenys olsoni
Strumigenys erynnes
90 Strumigenys ludovici
75 Strumigenys chyzeri
Strumigenys emmae
100 Strumigenys godeffroyi
77 Strumigenys rogeri
Strumigenys coveri
Strumigenys dicomas
)LJXUH6WJ

Apterogyna ZA01
Amblyopone australis
Proceratium stictum
99 Stigmatomma pallipes
91 Platythyrea mocquerysi
100 Paraponera clavata
100 Ankylomyrma coronacantha
Tatuidris WDWXVLD
100 Tatuidris tatusiaB
Lasius californicus
100 Myrmelachista flavocotea
Acanthoponera minor
64 Rhytidoponera chalybaea
62 Aneuretus simoni
100 Dolichoderus pustulatus
100 Tetraponera rufonigra
Myrmecia pyriformis
Nothomyrmecia macrops
Acanthognathus ocellatus
Acanthomyrmex ferox
Adelomyrmex SC01
Adlerzia froggatti
Atopomyrmex mocquerysi
Bariamyrma hispidula
Basiceros manni
Blepharidatta conops
Calyptomyrmex kaurus
Carebarella PA01
Cryptomyrmex boltoni
Cyphoidris exalta
Dacatria templaris
Dacetinops ignotus cf
Daceton armigerum
Diaphoromyrma sofiae
Dicroaspis KM01
Diplomorium longipenne
Dolopomyrmex pilatus
Eutetramorium mocquerysi
100 Gauromyrmex acanthinus
Lachnomyrmex scrobiculatus
Mayriella ebbei
Megalomyrmex silvestrii
Melissotarsus insularis
Monomorium denticulatum
Monomorium hanneli
Monomorium sechellense
Myrmicine genus26 PH02
Myrmicocrypta infuscata cf
Ocymyrmex fortior cf
Orectognathus versicolor
Oxyepoecus vezenyii
Paratopula TH01
Podomyrma silvicola
Proatta butteli
Rhopalomastix rothneyi
Rogeria nevadensis
Rogeria stigmatica
Solenopsis papuana
Stegomyrmex manni
Tetheamyrma subspongia
Tyrannomyrmex legatus
Vollenhovia emeryi
81 Xenomyrmex floridanus
Adelomyrmex paratristani
100 Baracidris sitra
Allomerus octoarticulatus
72 Wasmannia auropunctata
Austromorium flavigaster
90 58 Monomorium antarcticum
Carebara urichi
100 &DUHEDUDDIILQLV
Cataulacus ebrardi
60 Cataulacus hispidulus
Lophomyrmex ambiguus
100 Recurvidris TH01
Meranoplus castaneus
80 Meranoplus radamae
Monomorium brocha
54 Monomorium latinode
Monomorium ferodens
100 Monomorium fisheri
Royidris notorthotenes
100 Royidris longiseta
Myrmicaria carinata
100 Myrmicaria exigua
Malagidris jugum
93 MDODJLGULVEHOWL
Phalacromyrmex fugax
51 Pilotrochus besmerus
Apterostigma auriculatum
Microdaceton tibialis
90 Mycocepurus goeldii
100 Carebara nosindambo
Carebara alperti
100 Carebara AU01
100 Procryptocerus scabriusculus
Cephalotes atratus
74 Cephalotes texanus
99 Huberia brounii
Myrmisaraka brevis
100 Vitsika suspicax
100 Liomyrmex gestroi
Metapone madagascarica
100 Metapone PG01
98 Monomorium robustior
Monomorium criniceps
100 Monomorium destructor
53 Myrmecina graminicola
Myrmecina americana
100 Myrmecina PG01
74 Terataner bottegoi
Terataner MG02
Terataner MG27
100 Ancyridris polyrhachioides nr
Lordomyrma bhutanensis cf
Lordomyrma desupra
100 Propodilobus pingorum
91 Anillomyrma AFRC TZ 01
60 Anillomyrma decamera
Kempfidris inusualis
67 Tropidomyrmex elianae
100 Kartidris sparsipila
100 Cardiocondyla mauritanica
Cardiocondyla MY01
50 Cardiocondyla thoracica
64 Carebara vidua
100 65 Mesostruma browni
Colobostruma unicorna
Epopostruma monstrosa
98 Chalepoxenus kutteri
Myrmoxenus bernardi
Temnothorax poeyi
71 Temnothorax rugatulus
100 Vombisidris bilongrudi
Formicoxenus diversipilosus
Harpagoxenus sublaevis
100 Leptothorax muscorum complex
100 Ochetomyrmex neopolitus
100 Ochetomyrmex semipolitus
Tranopelta subterranea
97 Tranopelta gilva
94 Romblonella scrobifera
100 Poecilomyrma myrmecodiae
Rotastruma recava
73 Rotastruma stenoceps
Crematogaster acuta
100 Crematogaster sordidula
97 100 Crematogaster modiglianii
Crematogaster adrepens
100 Crematogaster emeryana
Dilobocondyla borneensis cf
59 Secostruma MY01
62 100 Perissomyrmex snyderi
Pristomyrmex orbiceps
100 Pristomyrmex punctatus
Nesomyrmex echinatinodis
100 Nesomyrmex wilda
100 Nesomyrmex madecassus
Nesomyrmex MG01
96 Nesomyrmex MG08
91 97 Eurhopalothrix australis
Eurhopalothrix gravis
76 Talaridris mandibularis
100 Octostruma EC01
Protalaridris armata
Rhopalothrix isthmica
100 88 Monomorium pharaonis
95 Monomorium ergatogyna
75 Monomorium fieldi nr
53 Monomorium madecassum
Monomorium exiguum
69 Monomorium MG04
99 83 Solenopsis mameti
Solenopsis fugax
71 Solenopsis ZA02
62 Solenopsis xyloni
Solenopsis molesta
Solenopsis succinea
Kalathomyrmex emeryi
98 Mycetarotes acutus
77 Mycetosoritis hartmanni
Cyphomyrmex cornutus
100 Cyphomyrmex longiscapus cf
100 69 Acromyrmex versicolor
Atta texana
Trachymyrmex arizonensis
92 Pheidole fimbriata
100 Pheidole rhea
100 Pheidole longispinosa
Anisopheidole antipodum
91 Pheidole hyatti
Pheidole lucioccipitalis
73 Pheidole pallidula
Machomyrma dispar
Pheidole clydei
Tetramorium MG125
100 Rhoptromyrmex transversinodis
Tetramorium CF01
93 56 Anergates atratulus
Strongylognathus testaceus
Teleutomyrmex kutteri
75 Tetramorium caespitum cf
59 Tetramorium taylori
81 Tetramorium spinosum
69 7HWUDPRULXPYHQDWRU
Tetramorium severini
Tetramorium validiusculum
97 Lenomyrmex colwelli
88 Strumigenys ambatrix
100 100 Strumigenys nitens
Strumigenys gundlachi
Strumigenys biolleyi
99 Strumigenys exiguaevitae
Strumigenys maxillaris
Strumigenys olsoni
99 95 Strumigenys rogeri
Strumigenys erynnes
99 Strumigenys ludovici
Strumigenys coveri
Strumigenys dicomas
91 Strumigenys ocypete
99 Strumigenys membranifera
58 Strumigenys chiricahua
Strumigenys chyzeri
Strumigenys emmae
Strumigenys godeffroyi
100 Hylomyrma blandiens
Pogonomyrmex angustus
86 94 Pogonomyrmex striatinodus
66 100 Pogonomyrmex imberbiculus
Pogonomyrmex subdentatus
Pogonomyrmex vermiculatus
100 Manica bradleyi
Manica rubida
100 Myrmica kotokui
57 74 Myrmica rugiventris
Myrmica striolagaster
100 Myrmica tahoensis
Aphaenogaster pythia
79 Aphaenogaster swammerdami
100 Aphaenogaster araneoides
Aphaenogaster albisetosa
100 100 Messor andrei
97 Aphaenogaster occidentalis
Messor denticornis
100 Messor wasmanni
Goniomma blanci
84 Oxyopomyrmex santschii
97 Stenamma dyscheres
Stenamma expolitum
Stenamma striatulum
)LJXUH6()D)

Apterogyna ZA01
Acanthoponera minor
Amblyopone australis
55 Paraponera clavata
Platythyrea mocquerysi
83 Rhytidoponera chalybaea
Aneuretus simoni
91 100 Dolichoderus pustulatus
81 Lasius californicus
100 Myrmelachista flavocotea
Myrmecia pyriformis
53 Nothomyrmecia macrops
86 Proceratium stictum
54 Stigmatomma pallipes
94 Tetraponera rufonigra
100 Ankylomyrma coronacantha
Tatuidris WDWXVLD
100 Tatuidris tatusiaB
54 Pogonomyrmex angustus
81 Hylomyrma blandiens
78 Pogonomyrmex striatinodus
91 84 Pogonomyrmex subdentatus
Pogonomyrmex imberbiculus
99 Pogonomyrmex vermiculatus
Manica bradleyi
99 Manica rubida
100 Myrmica kotokui
Myrmica rugiventris
Myrmica striolagaster
Myrmica tahoensis
Acanthognathus ocellatus
Apterostigma auriculatum
Atopomyrmex mocquerysi
Basiceros manni
Dacatria templaris
Dacetinops ignotus cf
Daceton armigerum
Diaphoromyrma sofiae
Diplomorium longipenne
Dolopomyrmex pilatus
Kartidris sparsipila
Lachnomyrmex scrobiculatus
Lenomyrmex colwelli
Mycocepurus goeldii
Myrmicocrypta infuscata cf
Octostruma EC01
Ocymyrmex fortior cf
Paratopula TH01
Procryptocerus scabriusculus
Protalaridris armata
Rhopalothrix isthmica
Stegomyrmex manni
Tetheamyrma subspongia
Vombisidris bilongrudi
54 Xenomyrmex floridanus
Calyptomyrmex kaurus
100 Cyphoidris exalta
Cataulacus ebrardi
97 Cataulacus hispidulus
Cephalotes atratus
100 Cephalotes texanus
Dicroaspis KM01
65 Proatta butteli
Melissotarsus insularis
55 Rhopalomastix rothneyi
Myrmicine genus26 PH02
99 Vollenhovia emeryi
Phalacromyrmex fugax
100 Pilotrochus besmerus
Pristomyrmex orbiceps
52 Pristomyrmex punctatus
100 Perissomyrmex snyderi
Dilobocondyla borneensis cf
76 98 Secostruma MY01
100 Eurhopalothrix australis
Eurhopalothrix gravis
100 Talaridris mandibularis
68 Harpagoxenus sublaevis
Formicoxenus diversipilosus
Leptothorax muscorum complex
57 Blepharidatta conops
100 Microdaceton tibialis
Allomerus octoarticulatus
99 Wasmannia auropunctata
77 Orectognathus versicolor
54 Colobostruma unicorna
Epopostruma monstrosa
100 Mesostruma browni
68 Ochetomyrmex neopolitus
100 Ochetomyrmex semipolitus
Tranopelta subterranea
96 Tranopelta gilva
94 Romblonella scrobifera
100 Poecilomyrma myrmecodiae
Rotastruma recava
100 Rotastruma stenoceps
Meranoplus castaneus
65 Meranoplus radamae
100 100 Cardiocondyla mauritanica
Cardiocondyla MY01
100 Cardiocondyla thoracica
51 Crematogaster sordidula
100 Crematogaster acuta
100 Crematogaster modiglianii
Crematogaster adrepens
100 Crematogaster emeryana
Nesomyrmex echinatinodis
97 Nesomyrmex wilda
98 88 Nesomyrmex madecassus
Nesomyrmex MG01
Nesomyrmex MG08
75 79 Lophomyrmex ambiguus
Adlerzia froggatti
100 Recurvidris TH01
91 Terataner bottegoi
Terataner MG02
Terataner MG27
99 99 Carebara vidua
100 Carebara alperti
54 Carebara AU01
100 Carebara nosindambo
Carebara urichi
&DUHEDUDDIILQLV
78 Kalathomyrmex emeryi
77 Mycetarotes acutus
99 Mycetosoritis hartmanni
76 Cyphomyrmex longiscapus cf
100 Cyphomyrmex cornutus
79 Trachymyrmex arizonensis
Acromyrmex versicolor
Atta texana
72 100 Gauromyrmex acanthinus
92 100 Myrmecina PG01
Myrmecina americana
53 Myrmecina graminicola
76 Temnothorax poeyi
99 Temnothorax rugatulus
Chalepoxenus kutteri
Myrmoxenus bernardi
99 Pheidole fimbriata
99 86 Pheidole rhea
Anisopheidole antipodum
100 Pheidole hyatti
52 Machomyrma dispar
Pheidole clydei
60 Pheidole longispinosa
Pheidole lucioccipitalis
99 Pheidole pallidula
98 Stenamma expolitum
Stenamma dyscheres
80 100 Stenamma striatulum
Aphaenogaster albisetosa
58 100 Messor andrei
Aphaenogaster pythia
60 Aphaenogaster swammerdami
93 100 Aphaenogaster araneoides
Goniomma blanci
99 Oxyopomyrmex santschii
100 Aphaenogaster occidentalis
Messor denticornis
100 Messor wasmanni
Malagidris jugum
69 Malagidris belti
51 100 Eutetramorium mocquerysi
Myrmisaraka brevis
Vitsika suspicax
69 97 Huberia brounii
100 Podomyrma silvicola
100 Liomyrmex gestroi
Metapone madagascarica
100 Metapone PG01
Royidris notorthotenes
73 Royidris longiseta
100 97 Monomorium robustior
Monomorium criniceps
Monomorium destructor
Acanthomyrmex ferox
100 Mayriella ebbei
80 Ancyridris polyrhachioides nr
Lordomyrma bhutanensis cf
60 Lordomyrma desupra
100 Propodilobus pingorum
7HWUDPRULXPYHQDWRU
100 Tetramorium validiusculum
Tetramorium CF01
100 100 Tetramorium taylori
78 Tetramorium caespitum cf
92 Strongylognathus testaceus
Anergates atratulus
Teleutomyrmex kutteri
69 Rhoptromyrmex transversinodis
51 Tetramorium MG125
Tetramorium severini
Tetramorium spinosum
97 Strumigenys ambatrix
98 Strumigenys nitens
96 Strumigenys gundlachi
100 Strumigenys biolleyi
Strumigenys erynnes
100 Strumigenys ludovici
100 64 Strumigenys rogeri
Strumigenys coveri
Strumigenys dicomas
96 Strumigenys maxillaris
67 96 Strumigenys olsoni
54 Strumigenys membranifera
63 Strumigenys ocypete
70 Strumigenys chiricahua
99 Strumigenys exiguaevitae
98 Strumigenys godeffroyi
Strumigenys chyzeri
Strumigenys emmae
90 Rogeria nevadensis
Bariamyrma hispidula
100 Rogeria stigmatica
Carebarella PA01
99 Solenopsis papuana
100 100 Solenopsis mameti
Solenopsis fugax
97 Solenopsis ZA02
62 73 Solenopsis molesta
Solenopsis succinea
Solenopsis xyloni
Megalomyrmex silvestrii
Monomorium brocha
Monomorium hanneli
100 Tyrannomyrmex legatus
Anillomyrma AFRC TZ 01
100 Anillomyrma decamera
Myrmicaria carinata
64 88 Myrmicaria exigua
53 Monomorium denticulatum
Austromorium flavigaster
76 Monomorium antarcticum
63 Adelomyrmex paratristani
100 Oxyepoecus vezenyii
Kempfidris inusualis
Tropidomyrmex elianae
Baracidris sitra
57 51 Monomorium sechellense
Adelomyrmex SC01
Cryptomyrmex boltoni
75 Monomorium ferodens
87 Monomorium fisheri
77 Monomorium latinode
Monomorium madecassum
85 Monomorium MG04
69 Monomorium pharaonis
60 Monomorium ergatogyna
Monomorium exiguum
Monomorium fieldi nr
)LJXUH6&$'

Apterogyna ZA01
100 Platythyrea mocquerysi
Proceratium stictum
100 Stigmatomma pallipes
65 100 Ankylomyrma coronacantha
Tatuidris WDWXVLD
93 Tatuidris tatusiaB
Amblyopone australis
65 Paraponera clavata
69 Tetraponera rufonigra
57 Acanthoponera minor
55 87 Rhytidoponera chalybaea
Aneuretus simoni
100 Dolichoderus pustulatus
94 Lasius californicus
100 Myrmelachista flavocotea
Myrmecia pyriformis
65 100
Nothomyrmecia macrops
68 Pogonomyrmex vermiculatus
Hylomyrma blandiens
100 Pogonomyrmex angustus
100 100 100 Pogonomyrmex striatinodus
Pogonomyrmex imberbiculus
100 Pogonomyrmex subdentatus
Manica bradleyi
100 Manica rubida
100 96 Myrmica kotokui
67 91 Myrmica rugiventris
Myrmica striolagaster
100 Myrmica tahoensis
Aphaenogaster albisetosa
100 100 Messor andrei
Stenamma dyscheres
Stenamma expolitum
98 98 Stenamma striatulum
Aphaenogaster araneoides
100 100 Aphaenogaster occidentalis
93 Messor denticornis
100 Messor wasmanni
91 Aphaenogaster pythia
100 Aphaenogaster swammerdami
Goniomma blanci
Oxyopomyrmex santschii
92 Stegomyrmex manni
70 Dolopomyrmex pilatus
72 Rogeria stigmatica
Bariamyrma hispidula
100 100 Rogeria nevadensis
Monomorium denticulatum
93 Oxyepoecus vezenyii
66 95 Megalomyrmex silvestrii
Austromorium flavigaster
100 87 Monomorium antarcticum
76 Monomorium brocha
Monomorium latinode
100 Tyrannomyrmex legatus
Kempfidris inusualis
94 100 Tropidomyrmex elianae
Carebarella PA01
100 98 Solenopsis papuana
100 100 Solenopsis xyloni
Solenopsis fugax
61 Solenopsis ZA02
100 Solenopsis mameti
Solenopsis molesta
92 Solenopsis succinea
Monomorium ferodens
Monomorium fisheri
100 Monomorium hanneli
Myrmicaria carinata
66 Myrmicaria exigua
100 100 Monomorium sechellense
Anillomyrma AFRC TZ 01
Anillomyrma decamera
99 Adelomyrmex paratristani
57 Baracidris sitra
Adelomyrmex SC01
Cryptomyrmex boltoni
100 Monomorium pharaonis
57 78 Monomorium madecassum
89 Monomorium ergatogyna
100 Monomorium fieldi nr
Monomorium exiguum
100 Monomorium MG04
100 Orectognathus versicolor
77 100 Colobostruma unicorna
Epopostruma monstrosa
99 Mesostruma browni
99 Ochetomyrmex neopolitus
100 Ochetomyrmex semipolitus
Tranopelta subterranea
58 Tranopelta gilva
Acanthognathus ocellatus
Diaphoromyrma sofiae
72 Lachnomyrmex scrobiculatus
Lenomyrmex colwelli
Microdaceton tibialis
Apterostigma auriculatum
91 74 99 Daceton armigerum
Mycocepurus goeldii
Myrmicocrypta infuscata cf
82 61 Kalathomyrmex emeryi
100 Mycetarotes acutus
100 Mycetosoritis hartmanni
98 63 Cyphomyrmex cornutus
100 Cyphomyrmex longiscapus cf
84 Trachymyrmex arizonensis
Acromyrmex versicolor
100 Atta texana
100 Blepharidatta conops
Allomerus octoarticulatus
100 Wasmannia auropunctata
100 Procryptocerus scabriusculus
92 Cephalotes atratus
Cephalotes texanus
100 Pheidole fimbriata
97 Pheidole rhea
100 100 Pheidole clydei
80 66 69 Anisopheidole antipodum
Machomyrma dispar
80 Pheidole longispinosa
57 Pheidole hyatti
Pheidole lucioccipitalis
Pheidole pallidula
99 100 Basiceros manni
Protalaridris armata
99 Rhopalothrix isthmica
99 Octostruma EC01
99 100 Eurhopalothrix australis
Eurhopalothrix gravis
100 Talaridris mandibularis
Phalacromyrmex fugax
98 Pilotrochus besmerus
Strumigenys ambatrix
100 100 100 Strumigenys nitens
Strumigenys gundlachi
Strumigenys biolleyi
99 Strumigenys exiguaevitae
Strumigenys maxillaris
100 Strumigenys olsoni
100 Strumigenys erynnes
100 Strumigenys ludovici
99 Strumigenys rogeri
Strumigenys coveri
Strumigenys dicomas
100 100 Strumigenys chiricahua
77 Strumigenys membranifera
99 Strumigenys ocypete
100 Strumigenys godeffroyi
59 Strumigenys chyzeri
Strumigenys emmae
Acanthomyrmex ferox
Atopomyrmex mocquerysi
Calyptomyrmex kaurus
Cyphoidris exalta
Dacetinops ignotus cf
Dicroaspis KM01
Gauromyrmex acanthinus
Kartidris sparsipila
Lophomyrmex ambiguus
Mayriella ebbei
Ocymyrmex fortior cf
Perissomyrmex snyderi
Recurvidris TH01
100 Tetheamyrma subspongia
Cataulacus ebrardi
53 Cataulacus hispidulus
Dacatria templaris
100 Diplomorium longipenne
Dilobocondyla borneensis cf
100 Secostruma MY01
Melissotarsus insularis
100 Rhopalomastix rothneyi
Meranoplus castaneus
99 Meranoplus radamae
Pristomyrmex orbiceps
64 Pristomyrmex punctatus
100 Adlerzia froggatti
Myrmicine genus26 PH02
91 Vollenhovia emeryi
Proatta butteli
Vombisidris bilongrudi
100 Xenomyrmex floridanus
54 Terataner bottegoi
Terataner MG02
100 Terataner MG27
69 Ancyridris polyrhachioides nr
73 Propodilobus pingorum
Lordomyrma bhutanensis cf
100 Lordomyrma desupra
Crematogaster acuta
100 Crematogaster sordidula
99 100 Crematogaster modiglianii
Crematogaster adrepens
95 100 Crematogaster emeryana
Nesomyrmex echinatinodis
100 Nesomyrmex wilda
100 88 Nesomyrmex MG08
Nesomyrmex madecassus
63 Nesomyrmex MG01
100 Paratopula TH01
69 Poecilomyrma myrmecodiae
100 Romblonella scrobifera
Rotastruma recava
100 Rotastruma stenoceps
67 59 Cardiocondyla mauritanica
Cardiocondyla MY01
100 Cardiocondyla thoracica
100 Myrmecina PG01
Myrmecina americana
Myrmecina graminicola
96 Carebara vidua
97 100 Carebara nosindambo
52 Carebara alperti
100 Carebara AU01
Carebara urichi
100 &DUHEDUDDIILQLV
100 76 Leptothorax muscorum complex
Formicoxenus diversipilosus
100 Harpagoxenus sublaevis
81 Temnothorax poeyi
99 Temnothorax rugatulus
Chalepoxenus kutteri
100 Myrmoxenus bernardi
Tetramorium CF01
100 86 Tetramorium taylori
100 Rhoptromyrmex transversinodis
82 100 Tetramorium MG125
Tetramorium severini
100 Tetramorium spinosum
7HWUDPRULXPYHQDWRU
73 Tetramorium validiusculum
100 100 Tetramorium caespitum cf
100 Strongylognathus testaceus
Anergates atratulus
100 Teleutomyrmex kutteri
100 Monomorium robustior
Monomorium criniceps
100 Monomorium destructor
100 Eutetramorium mocquerysi
Royidris notorthotenes
97 100 Royidris longiseta
80 Malagidris jugum
100 Malagidris belti
Myrmisaraka brevis
100 Vitsika suspicax
58 Podomyrma silvicola
100 Huberia brounii
100 Liomyrmex gestroi
Metapone madagascarica
Metapone PG01
Ward, P.S., Brady, S.G., Fisher, B.L., & Schultz, T.R. (2014) The evolution of myrmicine ants: phylogeny and biogeography of a hyperdiverse ant
clade (Hymenoptera: Formicidae). Systematic Entomology

Table S1. List of taxa sampled (n=251), voucher specimen codes, and GenBank accession numbers. Full collection data for all specimens can be
found by searching the voucher numbers on AntWeb (www.antweb.org).

Taxon Voucher 18S 28S Wg Abd-A LW Rh EF1aF1 EF1aF2 ArgK CAD Top1 Ubx
Acanthognathus ocellatus CASENT0007007 EF012824 EF012952 EF013660 EF013080 EF013532 EF013208 EF013370 KJ860918 KJ860677 KJ861519 KJ860248
Acanthomyrmex ferox CASENT0106243 KJ860060 KJ859872 KJ861760 KJ861149 KJ861335 KJ860489 KJ859686 KJ860919 KJ860678 KJ861520 KJ860249
Acromyrmex versicolor CASENT0106056 EF012826 EF012954 EF013662 EF013082 EF013534 EF013211 EF013373 KJ860920 KJ860679 KJ861521 KJ860250
Adelomyrmex paratristani CASENT0106201 KJ860061 KJ859873 KJ861761 KJ861150 KJ861336 KJ860490 KJ859687 KJ860921 KJ860680 KJ861522 KJ860251
Adelomyrmex SC01 CASENT0161591 KJ860062 KJ859874 KJ861762 KJ861151 KJ861337 KJ860491 KJ859688 KJ860922 KJ860681 KJ861523 KJ860252
Adlerzia froggatti CASENT0227591 KJ860063 KJ859875 KJ861763 KJ861152 KJ861338 KJ860492 KJ859689 KJ860923 KJ860682 KJ861524 KJ860253
Allomerus octoarticulatus CASENT0106199 KJ860064 KJ859876 KJ861764 KJ861153 KJ861339 KJ860493 KJ859690 KJ860924 KJ860683 KJ861525 KJ860254
Ancyridris polyrhachioides_nr CASENT0106303 KJ860065 KJ859877 KJ861765 KJ861154 KJ861340 KJ860494 KJ859691 KJ860925 KJ860684 KJ861526 KJ860255
Anergates atratulus CASENT0106265 KJ860066 KJ859878 KJ861766 KJ861155 KJ861341 KJ860495 KJ859692 KJ860926 KJ860685 KJ861527 KJ860256
Anillomyrma AFRC_TZ_01 CASENT0280595 KJ860067 KJ859879 KJ861767 KJ861156 KJ861342 KJ860496 KJ859693 KJ860927 KJ860686 KJ861528 KJ860257
Anillomyrma decamera CASENT0235147 KJ860068 KJ859880 KJ861768 KJ861157 KJ861343 KJ860497 KJ859694 KJ860928 KJ860687 KJ861529 KJ860258
Anisopheidole antipodum CASENT0106219 KJ860069 KJ859881 KJ861769 KJ861158 KJ861344 KJ860498 KJ859695 KJ860929 KJ860688 KJ861530 KJ860259
Aphaenogaster albisetosa CASENT0106040 EF012837 EF012965 EF013673 EF013093 EF013545 EF013224 EF013386 KJ860930 KJ860689 KJ861531 KJ860260
Aphaenogaster araneoides CASENT0106242 KJ860070 KJ859882 KJ861770 KJ861159 KJ861345 KJ860499 KJ859696 KJ860931 KJ860690 KJ861532 KJ860261
Aphaenogaster occidentalis CASENT0106090 AY867450 AY867466 AY867435 AY867482 AY867497 EF013225 EF013387 KJ860932 KJ860691 KJ861533 KJ860262
Aphaenogaster pythia CASENT0106131 KJ860071 KJ859883 KJ861771 KJ861160 KJ861346 KJ860500 KJ859697 KJ860933 KJ860692 KJ861534 KJ860263
Aphaenogaster swammerdami CASENT0498428 EF012838 EF012966 EF013674 EF013094 EF013546 EF013226 EF013388 KJ860934 KJ860693 KJ861535 KJ860264
Apterostigma auriculatum CASENT0010122 EF012841 EF012969 EF013677 EF013097 EF013549 EF013230 EF013392 KJ860935 KJ860694 KJ861536 KJ860265
Atopomyrmex mocquerysi CASENT0415562 KJ860072 KJ859884 KJ861772 KJ861161 KJ861347 KJ860501 KJ859698 KJ860936 KJ860695 KJ861537 KJ860266
Atta texana CASENT0106142 KJ860073 KJ859885 KJ861773 KJ861162 KJ861348 KJ860502 KJ859699 KJ860937 KJ860696 KJ861538 KJ860267
Austromorium flavigaster CASENT0106211 KJ860074 KJ859886 KJ861774 KJ861163 KJ861349 KJ860503 KJ859700 KJ860938 KJ860697 KJ861539 KJ860268
Baracidris sitra CASENT0006829 KJ860075 KJ859887 KJ861775 KJ861164 KJ861350 KJ860504 KJ859701 KJ860939 KJ860698 KJ861540 KJ860269
Bariamyrma hispidula INB0003662387 KJ860076 KJ859888 KJ861776 KJ861165 KJ861351 KJ860505 KJ859702 KJ860940 KJ860699 KJ861541 KJ860270
Basiceros manni CASENT0052769 EF012843 EF012971 EF013679 EF013099 EF013551 EF013232 EF013394 KJ860941 KJ860700 KJ861542 KJ860271
Blepharidatta conops CASENT0106248 KJ860077 KJ859889 KJ861777 KJ861166 KJ861352 KJ860506 KJ859703 KJ860942 KJ860701 KJ861543 KJ860272
Calyptomyrmex kaurus CASENT0417155 KJ860078 KJ859890 KJ861778 KJ861167 GU075846 KJ860507 KJ859704 GU085757 KJ860702 KJ861544 KJ860273
Cardiocondyla mauritanica CASENT0106115 EF012849 EF012977 EF013685 EF013105 EF013557 EF013239 EF013401 KJ860943 KJ860703 KJ861545 KJ860274
Cardiocondyla MY01 CASENT0106253 KJ860079 KJ859891 KJ861779 KJ861168 KJ861353 KJ860508 KJ859705 KJ860944 KJ860704 KJ861546 KJ860275
Cardiocondyla thoracica CASENT0106252 KJ860080 KJ859892 KJ861780 KJ861169 KJ861354 KJ860509 KJ859706 KJ860945 KJ860705 KJ861547 KJ860276
Carebara affinis CASENT0106016 EF012909 EF013037 EF013745 EF013165 EF013617 EF013319 EF013481 KJ860946 KJ860706 KJ861548 KJ860277
Carebara alperti CASENT0106280 KJ860081 KJ859893 KJ861781 KJ861170 KJ861355 KJ860510 KJ859707 KJ860947 KJ860707 KJ861549 KJ860278
Carebara AU01 CASENT0106279 KJ860082 KJ859894 KJ861782 KJ861171 KJ861356 KJ860511 KJ859708 KJ860948 KJ860708 KJ861550 KJ860279
Carebara nosindambo CASENT0494760 KJ860083 KJ859895 KJ861783 KJ861172 KJ861357 KJ860512 KJ859709 KJ860949 KJ860709 KJ861551 KJ860280
Carebara urichi CASENT0106295 KJ860084 KJ859896 KJ861784 KJ861173 KJ861358 KJ860513 KJ859710 KJ860950 KJ860710 KJ861552 KJ860281
Carebara vidua CASENT0260120 KJ860085 KJ859897 KJ861785 KJ861174 KJ861359 KJ860514 KJ859711 KJ860951 KJ860711 KJ861553 KJ860282
Carebarella PA01 CASENT0235223 KJ860086 KJ859898 KJ861786 KJ861175 KJ861360 KJ860515 KJ859712 KJ860952 KJ860712 KJ861554 KJ860283
Cataulacus ebrardi CASENT0487553 EF012850 EF012978 EF013686 EF013106 EF013558 EF013240 EF013402 KJ860953 KJ860713 KJ861555 KJ860284
Cataulacus hispidulus CASENT0106277 KJ860087 KJ859899 KJ861787 KJ861176 KJ861361 KJ860516 KJ859713 KJ860954 KJ860714 KJ861556 KJ860285
Cephalotes atratus CASENT0106296 KJ860088 KJ859900 KJ861788 KJ861177 KJ861362 KJ860517 KJ859714 KJ860955 KJ860715 KJ861557 KJ860286
Cephalotes texanus CASENT0106141 KJ860089 KJ859901 KJ861789 KJ861178 KJ861363 KJ860518 KJ859715 KJ860956 KJ860716 KJ861558 KJ860287
Chalepoxenus kutteri CASENT0106300 KJ860090 KJ859902 KJ861790 KJ861179 KJ861364 KJ860519 KJ859716 KJ860957 KJ860717 KJ861559 KJ860288
Colobostruma unicorna CASENT0106222 KJ860091 KJ859903 KJ861791 KJ861180 KJ861365 KJ860520 KJ859717 KJ860958 KJ860718 KJ861560 KJ860289
Crematogaster acuta CASENT0106245 KJ860092 KJ859904 KJ861792 KJ861181 KJ861366 KJ860521 KJ859718 KJ860959 KJ860719 KJ861561 KJ860290
Crematogaster emeryana CASENT0106034 EF012856 EF012984 EF013692 EF013112 EF013564 EF013249 EF013411 KJ860960 KJ860720 KJ861562 KJ860291

1
Crematogaster kelleri CASENT0498885 KJ860093 KJ859905 KJ861793 KJ861182 KJ861367 KJ860522 KJ859719 KJ860961 KJ860721 KJ861563 KJ860292
Crematogaster modiglianii CASENT0106281 KJ860094 KJ859906 KJ861794 KJ861183 KJ861368 KJ860523 KJ859720 KJ860962 KJ860722 KJ861564 KJ860293
Crematogaster sordidula CASENT0106292 KJ860095 KJ859907 KJ861795 KJ861184 KJ861369 KJ860524 KJ859721 KJ860963 KJ860723 KJ861565 KJ860294
Cryptomyrmex boltoni CASENT0260404 KJ860096 KJ859908 KJ861796 KJ861185 KJ861370 KJ860525 KJ859722 KJ860964 KJ860724 KJ861566 KJ860295
Cyphoidris exalta CASENT0405993 KJ860097 KJ859909 KJ861797 GQ410990 GQ411015 KJ860526 GQ410981 KJ860965 KJ860725 KJ861567 KJ860296
Cyphomyrmex cornutus CASENT0052778 KJ860098 KJ859910 KJ861798 KJ861186 KJ861371 KJ860527 KJ859723 KJ860966 KJ860726 KJ861568 KJ860297
Cyphomyrmex longiscapus_cf CASENT0052777 KJ860099 KJ859911 KJ861799 KJ861187 KJ861372 KJ860528 KJ859724 KJ860967 KJ860727 KJ861569 KJ860298
Dacatria templaris CASENT0235148 KJ860100 KJ859912 KJ861800 KJ861188 KJ861373 KJ860529 KJ859725 KJ860968 KJ860728 KJ861570 KJ860299
Dacetinops ignotus_cf CASENT0215429 KJ860101 KJ859913 KJ861801 KJ861189 KJ861374 KJ860530 KJ859726 KJ860969 KJ860729 KJ861571 KJ860300
Daceton armigerum CASENT0010124 EF012857 EF012985 EF013693 EF013113 EF013565 EF013251 EF013413 KJ860970 KJ860730 KJ861572 KJ860301
Diaphoromyrma sofiae CASENT0178237 KJ860102 KJ859914 KJ861802 KJ861190 KJ861375 KJ860531 KJ859727 KJ860971 KJ860731 KJ861573 KJ860302
Dicroaspis KM01 CASENT0145951 KJ860103 KJ859915 KJ861803 KJ861191 KJ861376 KJ860532 KJ859728 KJ860972 KJ860732 KJ861574 KJ860303
Dilobocondyla borneensis_cf CASENT0106130 KJ860104 KJ859916 KJ861804 KJ861192 KJ861377 KJ860533 KJ859729 KJ860973 KJ860733 KJ861575 KJ860304
Diplomorium longipenne CASENT0217046 KJ860105 KJ859917 KJ861805 KJ861193 KJ861378 KJ860534 KJ859730 KJ860974 KJ860734 KJ861576 KJ860305
Dolopomyrmex pilatus CASENT0106200 KJ860106 KJ859918 KJ861806 KJ861194 KJ861379 KJ860535 KJ859731 KJ860975 KJ860735 KJ861577 KJ860306
Epopostruma monstrosa CASENT0235851 KJ860107 KJ859919 KJ861807 KJ861195 KJ861380 KJ860536 KJ859732 KJ860976 KJ860736 KJ861578 KJ860307
Eurhopalothrix australis CASENT0106270 KJ860108 KJ859920 KJ861808 KJ861196 KJ861381 KJ860537 KJ859733 KJ860977 KJ860737 KJ861579 KJ860308
Eurhopalothrix gravis CASENT0107554 EF012864 EF012992 EF013700 EF013120 EF013572 EF013260 EF013422 KJ860978 KJ860738 KJ861580 KJ860309
Eutetramorium mocquerysi CASENT0077435 EF012865 EF012993 EF013701 EF013121 EF013573 EF013261 EF013423 KJ860979 KJ860739 KJ861581 KJ860310
Formicoxenus diversipilosus CASENT0106244 KJ860109 KJ859921 KJ861809 KJ861197 KJ861382 KJ860538 KJ859734 KJ860980 KJ860740 KJ861582 KJ860311
Gauromyrmex acanthinus CASENT0126225 KJ860110 KJ859922 KJ861810 KJ861198 KJ861383 KJ860539 KJ859735 KJ860981 KJ860741 KJ861583 KJ860312
Goniomma blanci CASENT0106230 KJ860111 KJ859923 KJ861811 KJ861199 KJ861384 KJ860540 KJ859736 KJ860982 KJ860742 KJ861584 KJ860313
Harpagoxenus sublaevis CASENT0106289 KJ860112 KJ859924 KJ861812 KJ861200 KJ861385 KJ860541 KJ859737 KJ860983 KJ860743 KJ861585 KJ860314
Huberia brounii CASENT0106136 KJ860113 KJ859925 KJ861813 KJ861201 KJ861386 KJ860542 KJ859738 KJ860984 KJ860744 KJ861586 KJ860315
Hylomyrma blandiens CASENT0106124 KJ860114 KJ859926 KJ861814 KJ861202 KJ861387 KJ860543 KJ859739 KJ860985 KJ860745 KJ861587 KJ860316
Kalathomyrmex emeryi CASENT0179595 KJ860115 KJ859927 KJ861815 KJ861203 KJ861388 KJ860544 KJ859740 KJ860986 KJ860746 KJ861588 KJ860317
Kartidris sparsipila CASENT0235130 KJ860116 KJ859928 KJ861816 KJ861204 KJ861389 KJ860545 KJ859741 KJ860987 KJ860747 KJ861589 KJ860318
Kempfidris inusualis CASENT0217050 KJ860117 KJ859929 KJ861817 KJ861205 KJ861390 KJ860546 KJ859742 KJ860988 KJ860748 KJ861590 KJ860319
Lachnomyrmex scrobiculatus CASENT0106202 KJ860118 KJ859930 KJ861818 KJ861206 KJ861391 KJ860547 KJ859743 KJ860989 KJ860749 KJ861591 KJ860320
Lenomyrmex colwelli CASENT0106210 KJ860119 KJ859931 KJ861819 KJ861207 KJ861392 KJ860548 KJ859744 KJ860990 KJ860750 KJ861592 KJ860321
Leptothorax muscorum_complex CASENT0106029 EF012874 EF013002 EF013710 EF013130 EF013582 EF013276 EF013438 KJ860991 KJ860751 KJ861593 KJ860322
Liomyrmex gestroi CASENT0106198 KJ860120 KJ859932 KJ861820 KJ861208 KJ861393 KJ860549 KJ859745 KJ860992 KJ860752 KJ861594 KJ860323
Lophomyrmex ambiguus CASENT0106225 KJ860121 KJ859933 KJ861821 KJ861209 KJ861394 KJ860550 KJ859746 KJ860993 KJ860753 KJ861595 KJ860324
Lordomyrma bhutanensis_cf CASENT0106234 KJ860122 KJ859934 KJ861822 KJ861210 KJ861395 KJ860551 KJ859747 KJ860994 KJ860754 KJ861596 KJ860325
Lordomyrma desupra CASENT0106145 KJ860123 KJ859935 KJ861823 GQ410991 GQ411016 KJ860552 GQ410982 GU085766 KJ860755 KJ861597 KJ860326
Machomyrma dispar CASENT0106247 KJ860124 KJ859936 KJ861824 KJ861211 KJ861396 KJ860553 KJ859748 KJ860995 KJ860756 KJ861598 KJ860327
Malagidris belti CASENT0119117 KJ860125 KJ859937 KJ861825 KJ861212 KJ861397 KJ860554 KJ859749 KJ860996 KJ860757 KJ861599 KJ860328
Malagidris jugum CASENT0494278 KJ860126 KJ859938 KJ861826 KJ861213 KJ861398 KJ860555 KJ859750 KJ860997 KJ860758 KJ861600 KJ860329
Manica bradleyi CASENT0106022 EF012878 EF013006 EF013714 EF013134 EF013586 EF013281 EF013443 FJ939877 FJ939931 KJ523698 KJ523615
Manica rubida CASENT0106288 KJ860127 KJ859939 KJ861827 KJ861214 KJ861399 KJ860556 KJ859751 KJ860998 KJ860759 KJ861601 KJ860330
Mayriella ebbei CASENT0106116 EF012879 EF013007 EF013715 EF013135 EF013587 EF013282 EF013444 GU085788 KJ860760 KJ861602 KJ860331
Megalomyrmex silvestrii CASENT0106196 KJ860128 KJ859940 KJ861828 KJ861215 KJ861400 KJ860557 KJ859752 KJ860999 KJ860761 KJ861603 KJ860332
Melissotarsus insularis CASENT0498665 KJ860129 KJ859941 KJ861829 KJ861216 KJ861401 KJ860558 KJ859753 KJ861000 KJ860762 KJ861604 KJ860333
Meranoplus castaneus CASENT0106276 KJ860130 KJ859942 KJ861830 KJ861217 KJ861402 KJ860559 KJ859754 KJ861001 KJ860763 KJ861605 KJ860334
Meranoplus radamae CASENT0486686 EF012880 EF013008 EF013716 EF013136 EF013588 EF013283 EF013445 GU085789 KJ860764 KJ861606 KJ860335
Mesostruma browni QMT112783 KJ860131 KJ859943 KJ861831 KJ861218 KJ861403 KJ860560 KJ859755 KJ861002 KJ860765 KJ861607 KJ860336
Messor andrei CASENT0106051 EF012881 EF013009 EF013717 EF013137 EF013589 EF013284 EF013446 KJ861003 KJ860766 KJ861608 KJ860337
Messor denticornis CASENT0106118 EF012882 EF013010 EF013718 EF013138 EF013590 EF013285 EF013447 KJ861004 KJ860767 KJ861609 KJ860338
Messor wasmanni CASENT0106293 KJ860132 KJ859944 KJ861832 KJ861219 KJ861404 KJ860561 KJ859756 KJ861005 KJ860768 KJ861610 KJ860339
Metapone madagascarica CASENT0004528 EF012884 EF013012 EF013720 EF013140 EF013592 EF013287 EF013449 KJ861006 KJ860769 KJ861611 KJ860340
Metapone PG01 CASENT0106132 KJ860133 KJ859945 KJ861833 KJ861220 KJ861405 KJ860562 KJ859757 KJ861007 KJ860770 KJ861612 KJ860341
Microdaceton tibialis CASENT0402199 EF012885 EF013013 EF013721 EF013141 EF013593 EF013288 EF013450 KJ861008 KJ860771 KJ861613 KJ860342
Monomorium antarcticum CASENT0106264 KJ860134 KJ859946 KJ861834 KJ861221 KJ861406 KJ860563 KJ859758 KJ861009 KJ860772 KJ861614 KJ860343
Monomorium brocha CASENT0616568 KJ860135 KJ859947 KJ861835 KJ861222 KJ861407 KJ860564 KJ859759 KJ861010 KJ860773 KJ861615 KJ860344

2
Monomorium criniceps CASENT0106263 KJ860136 KJ859948 KJ861836 KJ861223 KJ861408 KJ860565 KJ859760 KJ861011 KJ860774 KJ861616 KJ860345
Monomorium denticulatum CASENT0106208 KJ860137 KJ859949 KJ861837 KJ861224 KJ861409 KJ860566 KJ859761 KJ861012 KJ860775 KJ861617 KJ860346
Monomorium destructor CASENT0147465 KJ860138 KJ859950 KJ861838 KJ861225 KJ861410 KJ860567 KJ859762 KJ861013 KJ860776 KJ861618 KJ860347
Monomorium ergatogyna CASENT0106030 EF012886 EF013014 EF013722 EF013142 EF013594 EF013290 EF013452 KJ861014 KJ860777 KJ861619 KJ860348
Monomorium exiguum CASENT0148842 KJ860139 KJ859951 KJ861839 KJ861226 KJ861411 KJ860568 KJ859763 KJ861015 KJ860778 KJ861620 KJ860349
Monomorium ferodens CASENT0478937 KJ860140 KJ859952 KJ861840 KJ861227 KJ861412 KJ860569 KJ859764 KJ861016 KJ860779 KJ861621 KJ860350
Monomorium fieldi_nr CASENT0106273 KJ860141 KJ859953 KJ861841 KJ861228 KJ861413 KJ860570 KJ859765 KJ861017 KJ860780 KJ861622 KJ860351
Monomorium fisheri CASENT0491610 KJ860142 KJ859954 KJ861842 KJ861229 KJ861414 KJ860571 KJ859766 KJ861018 KJ860781 KJ861623 KJ860352
Monomorium hanneli CASENT0151856 KJ860143 KJ859955 KJ861843 KJ861230 KJ861415 KJ860572 KJ859767 KJ861019 KJ860782 KJ861624 KJ860353
Monomorium latinode CASENT0146495 KJ860144 KJ859956 KJ861844 KJ861231 KJ861416 KJ860573 KJ859768 KJ861020 KJ860783 KJ861625 KJ860354
Monomorium madecassum CASENT0147108 KJ860145 KJ859957 KJ861845 KJ861232 KJ861417 KJ860574 KJ859769 KJ861021 KJ860784 KJ861626 KJ860355
Monomorium MG04 CASENT0006834 KJ860146 KJ859958 KJ861846 KJ861233 KJ861418 KJ860575 KJ859770 KJ861022 KJ860785 KJ861627 KJ860356
Monomorium pharaonis CASENT0137619 KJ860147 KJ859959 KJ861847 KJ861234 KJ861419 KJ860576 KJ859771 KJ861023 KJ860786 KJ861628 KJ860357
Monomorium robustior CASENT0147087 KJ860148 KJ859960 KJ861848 KJ861235 KJ861420 KJ860577 KJ859772 KJ861024 KJ860787 KJ861629 KJ860358
Monomorium sechellense CASENT0158893 KJ860149 KJ859961 KJ861849 KJ861236 KJ861421 KJ860578 KJ859773 KJ861025 KJ860788 KJ861630 KJ860359
Mycetarotes acutus USNMENT00304226 KJ860150 KJ859962 KJ861850 KJ861237 KJ861422 KJ860579 KJ859774 KJ861026 KJ860789 KJ861631 KJ860360
Mycetosoritis hartmanni USNMENT00445590 KJ860151 KJ859963 KJ861851 KJ861238 KJ861423 KJ860580 KJ859775 KJ861027 KJ860790 KJ861632 KJ860361
Mycocepurus goeldii USNMENT00538752 KJ860152 KJ859964 KJ861852 KJ861239 KJ861424 KJ860581 KJ859776 KJ861028 KJ860791 KJ861633 KJ860362
Myrmecina americana CASENT0106284 KJ860153 KJ859965 KJ861853 KJ861240 KJ861425 KJ860582 KJ859777 KJ861029 KJ860792 KJ861634 KJ860363
Myrmecina graminicola CASENT0106054 EF012887 EF013015 EF013723 EF013143 EF013595 EF013293 EF013455 GU085790 KJ860793 KJ861635 KJ860364
Myrmecina PG01 CASENT0106283 KJ860154 KJ859966 KJ861854 KJ861241 KJ861426 KJ860583 KJ859778 KJ861030 KJ860794 KJ861636 KJ860365
Myrmica kotokui CASENT0106286 KJ860155 KJ859967 KJ861855 KJ861242 KJ861427 KJ860584 KJ859779 KJ861031 KJ860795 KJ861637 KJ860366
Myrmica rugiventris CASENT0106212 KJ860156 KJ859968 KJ861856 KJ861243 KJ861428 KJ860585 KJ859780 KJ861032 KJ860796 KJ861638 KJ860367
Myrmica striolagaster CASENT0106036 EF012890 EF013018 EF013726 EF013146 EF013598 EF013296 EF013458 KJ861033 KJ860797 KJ861639 KJ860368
Myrmica tahoensis CASENT0106091 AY703495 AY703562 AY703629 AY703696 AY703763 EF013297 EF013459 GU085791 KJ860798 KJ861640 KJ860369
Myrmicaria carinata CASENT0106282 KJ860157 KJ859969 KJ861857 KJ861244 KJ861429 KJ860586 KJ859781 KJ861034 KJ860799 KJ861641 KJ860370
Myrmicaria exigua CASENT0403455 EF012891 EF013019 EF013727 EF013147 EF013599 EF013298 EF013460 KJ861035 KJ860800 KJ861642 KJ860371
Myrmicine_genus26 PH02 CASENT0106232 KJ860158 KJ859970 KJ861858 KJ861245 KJ861430 KJ860587 KJ859782 KJ861036 KJ860801 KJ861643 KJ860372
Myrmicocrypta infuscata_cf CASENT0010123 EF012892 EF013020 EF013728 EF013148 EF013600 EF013299 EF013461 KJ861037 KJ860802 KJ861644 KJ860373
Myrmisaraka brevis CASENT0494750 KJ860159 KJ859971 KJ861859 KJ861246 KJ861431 KJ860588 KJ859783 KJ861038 KJ860803 KJ861645 KJ860374
Myrmoxenus bernardi CASENT0106299 KJ860160 KJ859972 KJ861860 KJ861247 KJ861432 KJ860589 KJ859784 KJ861039 KJ860804 KJ861646 KJ860375
Nesomyrmex echinatinodis CASENT0106044 EF012895 EF013023 EF013731 EF013151 EF013603 EF013303 EF013465 KJ861040 KJ860805 KJ861647 KJ860376
Nesomyrmex madecassus CASENT0487130 KJ860161 KJ859973 KJ861861 KJ861248 KJ861433 KJ860590 KJ859785 KJ861041 KJ860806 KJ861648 KJ860377
Nesomyrmex MG01 CASENT0460666 KJ860162 KJ859974 KJ861862 KJ861249 KJ861434 KJ860591 KJ859786 KJ861042 KJ860807 KJ861649 KJ860378
Nesomyrmex MG08 CASENT0008660 KJ860163 KJ859975 KJ861863 KJ861250 KJ861435 KJ860592 KJ859787 KJ861043 KJ860808 KJ861650 KJ860379
Nesomyrmex wilda CASENT0106231 KJ860164 KJ859976 KJ861864 KJ861251 KJ861436 KJ860593 KJ859788 KJ861044 KJ860809 KJ861651 KJ860380
Ochetomyrmex neopolitus CASENT0106223 KJ860165 KJ859977 KJ861865 KJ861252 KJ861437 KJ860594 KJ859789 KJ861045 KJ860810 KJ861652 KJ860381
Ochetomyrmex semipolitus CASENT0179593 KJ860166 KJ859978 KJ861866 KJ861253 KJ861438 KJ860595 KJ859790 KJ861046 KJ860811 KJ861653 KJ860382
Octostruma EC01 CASENT0008663 KJ860167 KJ859979 KJ861867 KJ861254 KJ861439 KJ860596 KJ859791 KJ861047 KJ860812 KJ861654 KJ860383
Ocymyrmex fortior_cf CASENT0106140 KJ860168 KJ859980 KJ861868 KJ861255 KJ861440 KJ860597 KJ859792 KJ861048 KJ860813 KJ861655 KJ860384
Orectognathus versicolor CASENT0106002 EF012903 EF013031 EF013739 EF013159 EF013611 EF013312 EF013474 KJ861049 KJ860814 KJ861656 KJ860385
Oxyepoecus vezenyii CASENT0217325 KJ860169 KJ859981 KJ861869 KJ861256 KJ861441 KJ860598 KJ859793 KJ861050 KJ860815 KJ861657 KJ860386
Oxyopomyrmex santschii CASENT0106267 KJ860170 KJ859982 KJ861870 KJ861257 KJ861442 KJ860599 KJ859794 KJ861051 KJ860816 KJ861658 KJ860387
Paratopula TH01 CASENT0010838 KJ860171 KJ859983 KJ861871 KJ861258 KJ861443 KJ860600 KJ859795 KJ861052 KJ860817 KJ861659 KJ860388
Perissomyrmex snyderi CASENT0106226 KJ860172 KJ859984 KJ861872 KJ861259 KJ861444 KJ860601 KJ859796 KJ861053 KJ860818 KJ861660 KJ860389
Phalacromyrmex fugax CASENT0217029 KJ860173 KJ859985 KJ861873 KJ861260 KJ861445 KJ860602 KJ859797 KJ861054 KJ860819 KJ861661 KJ860390
Pheidole clydei CASENT0106117 EF012907 EF013035 EF013743 EF013163 EF013615 EF013317 EF013479 KJ861055 KJ860820 KJ861662 KJ860391
Pheidole fimbriata CASENT0235906 KJ860174 KJ859986 KJ861874 KJ861261 KJ861446 KJ860603 KJ859798 KJ861056 KJ860821 KJ861663 KJ860392
Pheidole hyatti CASENT0106046 EF012908 EF013036 EF013744 EF013164 EF013616 EF013318 EF013480 KJ861057 KJ860822 KJ861664 KJ860393
Pheidole longispinosa CASENT0494701 KJ860175 KJ859987 KJ861875 KJ861262 KJ861447 KJ860604 KJ859799 KJ861058 KJ860823 KJ861665 KJ860394
Pheidole lucioccipitalis CASENT0106275 KJ860176 KJ859988 KJ861876 KJ861263 KJ861448 KJ860605 KJ859800 KJ861059 KJ860824 KJ861666 KJ860395
Pheidole pallidula CASENT0106290 KJ860177 KJ859989 KJ861877 KJ861264 KJ861449 KJ860606 KJ859801 KJ861060 KJ860825 KJ861667 KJ860396
Pheidole rhea CASENT0106262 KJ860178 KJ859990 KJ861878 KJ861265 KJ861450 KJ860607 KJ859802 KJ861061 KJ860826 KJ861668 KJ860397
Pilotrochus besmerus CASENT0047617 EF012911 EF013039 EF013747 EF013167 EF013619 EF013321 EF013483 KJ861062 KJ860827 KJ861669 KJ860398

3
Podomyrma silvicola CASENT0106143 KJ860179 KJ859991 KJ861879 KJ861266 KJ861451 KJ860608 KJ859803 KJ861063 KJ860828 KJ861670 KJ860399
Poecilomyrma myrmecodiae CASENT0106144 KJ860180 KJ859992 KJ861880 KJ861267 KJ861452 KJ860609 KJ859804 KJ861064 KJ860829 KJ861671 KJ860400
Pogonomyrmex angustus CASENT0106125 KJ860181 KJ859993 KJ861881 KJ861268 KJ861453 KJ860610 KJ859805 KJ861065 KJ860830 KJ861672 KJ860401
Pogonomyrmex imberbiculus CASENT0005713 KJ860182 KJ859994 KJ861882 KJ861269 KJ861454 KJ860611 KJ859806 KJ861066 KJ860831 KJ861673 KJ860402
Pogonomyrmex striatinodus CASENT0179592 KJ860183 KJ859995 KJ861883 KJ861270 KJ861455 KJ860612 KJ859807 KJ861067 KJ860832 KJ861674 KJ860403
Pogonomyrmex subdentatus CASENT0106024 EF012914 EF013042 EF013750 EF013170 EF013622 EF013325 EF013487 KJ861068 KJ860833 KJ861675 KJ860404
Pogonomyrmex vermiculatus CASENT0106213 KJ860184 KJ859996 KJ861884 KJ861271 KJ861456 KJ860613 KJ859808 KJ861069 KJ860834 KJ861676 KJ860405
Pristomyrmex orbiceps CASENT0417737 KJ860185 KJ859997 KJ861885 KJ861272 KJ861457 KJ860614 KJ859809 KJ861070 KJ860835 KJ861677 KJ860406
Pristomyrmex punctatus CASENT0106274 KJ860186 KJ859998 KJ861886 KJ861273 KJ861458 KJ860615 KJ859810 KJ861071 KJ860836 KJ861678 KJ860407
Proatta butteli CASENT0007617 KJ860187 KJ859999 KJ861887 KJ861274 KJ861459 KJ860616 KJ859811 KJ861072 KJ860837 KJ861679 KJ860408
Procryptocerus scabriusculus CASENT0106043 EF012924 EF013052 EF013760 EF013180 EF013632 EF013336 EF013498 KJ861073 KJ860838 KJ861680 KJ860409
Propodilobus pingorum Antbase.net01463 KJ860188 KJ860000 KJ861888 KJ861275 KJ861460 KJ860617 KJ859812 KJ861074 KJ860839 KJ861681 KJ860410
Protalaridris armata CASENT0008665 KJ860189 KJ860001 KJ861889 KJ861276 KJ861461 KJ860618 KJ859813 KJ861075 KJ860840 KJ861682 KJ860411
Recurvidris TH01 CASENT0131659 KJ860190 KJ860002 KJ861890 KJ861277 KJ861462 KJ860619 KJ859814 KJ861076 KJ860841 KJ861683 KJ860412
Rhopalomastix rothneyi CASENT0106013 EF012929 EF013057 EF013765 EF013185 EF013637 EF013342 EF013504 KJ861077 KJ860842 KJ861684 KJ860413
Rhopalothrix isthmica CASENT0235905 KJ860191 KJ860003 KJ861891 KJ861278 KJ861463 KJ860620 KJ859815 KJ861078 KJ860843 KJ861685 KJ860414
Rhoptromyrmex transversinodis CASENT0106251 KJ860192 KJ860004 KJ861892 KJ861279 KJ861464 KJ860621 KJ859816 KJ861079 KJ860844 KJ861686 KJ860415
Rogeria nevadensis CASENT0106209 KJ860193 KJ860005 KJ861893 KJ861280 KJ861465 KJ860622 KJ859817 KJ861080 KJ860845 KJ861687 KJ860416
Rogeria stigmatica CASENT0106171 KJ860194 KJ860006 KJ861894 KJ861281 GU075845 KJ860623 KJ859818 GU085792 KJ860846 KJ861688 KJ860417
Romblonella scrobifera CASENT0106224 KJ860195 KJ860007 KJ861895 KJ861282 KJ861466 KJ860624 KJ859819 KJ861081 KJ860847 KJ861689 KJ860418
Rotastruma recava CASENT0106250 KJ860196 KJ860008 KJ861896 KJ861283 KJ861467 KJ860625 KJ859820 KJ861082 KJ860848 KJ861690 KJ860419
Rotastruma stenoceps CASENT0106285 KJ860197 KJ860009 KJ861897 KJ861284 KJ861468 KJ860626 KJ859821 KJ861083 KJ860849 KJ861691 KJ860420
Royidris longiseta CASENT0120396 KJ860198 KJ860010 KJ861898 KJ861285 KJ861469 KJ860627 KJ859822 KJ861084 KJ860850 KJ861692 KJ860421
Royidris notorthotenes CASENT0002259 KJ860199 KJ860011 KJ861899 KJ861286 KJ861470 KJ860628 KJ859823 KJ861085 KJ860851 KJ861693 KJ860422
Secostruma MY01 CASENT0217352 KJ860200 KJ860012 KJ861900 KJ861287 KJ861471 KJ860629 KJ859824 KJ861086 KJ860852 KJ861694 KJ860423
Solenopsis fugax CASENT0106291 KJ860201 KJ860013 KJ861901 KJ861288 KJ861472 KJ860630 KJ859825 KJ861087 KJ860853 KJ861695 KJ860424
Solenopsis mameti CASENT0006828 KJ860202 KJ860014 KJ861902 KJ861289 KJ861473 KJ860631 KJ859826 KJ861088 KJ860854 KJ861696 KJ860425
Solenopsis molesta CASENT0106027 EF012934 EF013062 EF013770 EF013190 EF013642 EF013348 EF013510 KJ861089 KJ860855 KJ861697 KJ860426
Solenopsis papuana CASENT0106272 KJ860203 KJ860015 KJ861903 KJ861290 KJ861474 KJ860632 KJ859827 KJ861090 KJ860856 KJ861698 KJ860427
Solenopsis succinea CASENT0106271 KJ860204 KJ860016 KJ861904 KJ861291 KJ861475 KJ860633 KJ859828 KJ861091 KJ860857 KJ861699 KJ860428
Solenopsis xyloni CASENT0106041 EF012935 EF013063 EF013771 EF013191 EF013643 EF013349 EF013511 KJ861092 KJ860858 KJ861700 KJ860429
Solenopsis ZA02 CASENT0217036 KJ860205 KJ860017 KJ861905 KJ861292 KJ861476 KJ860634 KJ859829 KJ861093 KJ860859 KJ861701 KJ860430
Stegomyrmex manni INB0003692959 KJ860206 KJ860018 KJ861906 KJ861293 KJ861477 KJ860635 KJ859830 KJ861094 KJ860860 KJ861702 KJ860431
Stenamma dyscheres CASENT0106023 EF012936 EF013064 EF013772 EF013192 EF013644 EF013351 EF013513 KJ861095 KJ860861 KJ861703 KJ860432
Stenamma expolitum CASENT0052774 KJ860207 KJ860019 KJ861907 KJ861294 KJ861478 KJ860636 KJ859831 GU085793 KJ860862 KJ861704 KJ860433
Stenamma striatulum CASENT0106294 KJ860208 KJ860020 KJ861908 KJ861295 KJ861479 KJ860637 KJ859832 KJ861096 KJ860863 KJ861705 KJ860434
Strongylognathus testaceus CASENT0106266 KJ860209 KJ860021 KJ861909 KJ861296 KJ861480 KJ860638 KJ859833 KJ861097 KJ860864 KJ861706 KJ860435
Strumigenys ambatrix CASENT0140727 KJ860210 KJ860022 KJ861910 KJ861297 KJ861481 KJ860639 KJ859834 KJ861098 KJ860865 KJ861707 KJ860436
Strumigenys biolleyi CASENT0106237 KJ860211 KJ860023 KJ861911 KJ861298 KJ861482 KJ860640 KJ859835 KJ861099 KJ860866 KJ861708 KJ860437
Strumigenys chiricahua CASENT0226156 KJ860212 KJ860024 KJ861912 KJ861299 KJ861483 KJ860641 KJ859836 KJ861100 KJ860867 KJ861709 KJ860438
Strumigenys chyzeri CASENT0106249 KJ860213 KJ860025 KJ861913 KJ861300 KJ861484 KJ860642 KJ859837 KJ861101 KJ860868 KJ861710 KJ860439
Strumigenys coveri CASENT0153050 KJ860214 KJ860026 KJ861914 KJ861301 KJ861485 KJ860643 KJ859838 KJ861102 KJ860869 KJ861711 KJ860440
Strumigenys dicomas CASENT0499800 EF012937 EF013065 EF013773 EF013193 EF013645 EF013352 EF013514 KJ861103 KJ860870 KJ861712 KJ860441
Strumigenys emmae CASENT0162633 KJ860215 KJ860027 KJ861915 KJ861302 KJ861486 KJ860644 KJ859839 KJ861104 KJ860871 KJ861713 KJ860442
Strumigenys erynnes CASENT0151597 KJ860216 KJ860028 KJ861916 KJ861303 KJ861487 KJ860645 KJ859840 KJ861105 KJ860872 KJ861714 KJ860443
Strumigenys exiguaevitae CASENT0456151 EF012928 EF013056 EF013764 EF013184 EF013636 EF013341 EF013503 KJ861106 KJ860873 KJ861715 KJ860444
Strumigenys godeffroyi CASENT0159907 KJ860217 KJ860029 KJ861917 KJ861304 KJ861488 KJ860646 KJ859841 KJ861107 KJ860874 KJ861716 KJ860445
Strumigenys gundlachi CASENT0106238 KJ860218 KJ860030 KJ861918 KJ861305 KJ861489 KJ860647 KJ859842 KJ861108 KJ860875 KJ861717 KJ860446
Strumigenys ludovici CASENT0152373 KJ860219 KJ860031 KJ861919 KJ861306 KJ861490 KJ860648 KJ859843 KJ861109 KJ860876 KJ861718 KJ860447
Strumigenys maxillaris CASENT0133614 KJ860220 KJ860032 KJ861920 KJ861307 KJ861491 KJ860649 KJ859844 KJ861110 KJ860877 KJ861719 KJ860448
Strumigenys membranifera CASENT0149215 KJ860221 KJ860033 KJ861921 KJ861308 KJ861492 KJ860650 KJ859845 KJ861111 KJ860878 KJ861720 KJ860449
Strumigenys nitens CASENT0106246 KJ860222 KJ860034 KJ861922 KJ861309 KJ861493 KJ860651 KJ859846 KJ861112 KJ860879 KJ861721 KJ860450
Strumigenys ocypete CASENT0217341 KJ860223 KJ860035 KJ861923 KJ861310 KJ861494 KJ860652 KJ859847 KJ861113 KJ860880 KJ861722 KJ860451
Strumigenys olsoni CASENT0141982 KJ860224 KJ860036 KJ861924 KJ861311 KJ861495 KJ860653 KJ859848 KJ861114 KJ860881 KJ861723 KJ860452

4
Strumigenys rogeri CASENT0158880 KJ860225 KJ860037 KJ861925 KJ861312 KJ861496 KJ860654 KJ859849 KJ861115 KJ860882 KJ861724 KJ860453
Talaridris mandibularis CASENT0235384 KJ860226 KJ860038 KJ861926 KJ861313 KJ861497 KJ860655 KJ859850 KJ861116 KJ860883 KJ861725 KJ860454
Teleutomyrmex kutteri CASENT0106301 KJ860227 KJ860039 KJ861927 KJ861314 KJ861498 KJ860656 KJ859851 KJ861117 KJ860884 KJ861726 KJ860455
Temnothorax poeyi CASENT0106241 KJ860228 KJ860040 KJ861928 KJ861315 KJ861499 KJ860657 KJ859852 KJ861118 KJ860885 KJ861727 KJ860456
Temnothorax rugatulus CASENT0106025 EF012942 EF013070 EF013778 EF013198 EF013650 EF013357 EF013519 GU085794 KJ860886 KJ861728 KJ860457
Terataner bottegoi CASENT0106298 KJ860229 KJ860041 KJ861929 KJ861316 KJ861500 KJ860658 KJ859853 KJ861119 KJ860887 KJ861729 KJ860458
Terataner MG02 CASENT0494349 EF012943 EF013071 EF013779 EF013199 EF013651 EF013358 EF013520 KJ861120 KJ860888 KJ861730 KJ860459
Terataner MG27 CASENT0494256 KJ860230 KJ860042 KJ861930 KJ861317 KJ861501 KJ860659 KJ859854 KJ861121 KJ860889 KJ861731 KJ860460
Tetheamyrma subspongia CASENT0106305 KJ860231 KJ860043 KJ861931 KJ861318 KJ861502 KJ860660 KJ859855 KJ861122 KJ860890 KJ861732 KJ860461
Tetramorium caespitum_cf CASENT0106026 EF012944 EF013072 EF013780 EF013200 EF013652 EF013359 EF013521 KJ861123 KJ860891 KJ861733 KJ860462
Tetramorium CF01 CASENT0006833 KJ860232 KJ860044 KJ861932 KJ861319 KJ861503 KJ860661 KJ859856 KJ861124 KJ860892 KJ861734 KJ860463
Tetramorium MG125 CASENT0048793 KJ860233 KJ860045 KJ861933 KJ861320 KJ861504 KJ860662 KJ859857 KJ861125 KJ860893 KJ861735 KJ860464
Tetramorium severini CASENT0494149 KJ860234 KJ860046 KJ861934 KJ861321 KJ861505 KJ860663 KJ859858 KJ861126 KJ860894 KJ861736 KJ860465
Tetramorium spinosum CASENT0106287 KJ860235 KJ860047 KJ861935 KJ861322 KJ861506 KJ860664 KJ859859 KJ861127 KJ860895 KJ861737 KJ860466
Tetramorium taylori CASENT0106297 KJ860236 KJ860048 KJ861936 KJ861323 KJ861507 KJ860665 KJ859860 KJ861128 KJ860896 KJ861738 KJ860467
Tetramorium validiusculum CASENT0106004 EF012945 EF013073 EF013781 EF013201 EF013653 EF013360 EF013522 KJ861129 KJ860897 KJ861739 KJ860468
Tetramorium venator CASENT0415619 KJ860237 KJ860049 KJ861937 KJ861324 KJ861508 KJ860666 KJ859861 KJ861130 KJ860898 KJ861740 KJ860469
Trachymyrmex arizonensis CASENT0106047 EF012947 EF013075 EF013783 EF013203 EF013655 EF013364 EF013526 KJ861131 KJ860899 KJ861741 KJ860470
Tranopelta gilva CASENT0106206 KJ860238 KJ860050 KJ861938 KJ861325 KJ861509 KJ860667 KJ859862 KJ861132 KJ860900 KJ861742 KJ860471
Tranopelta subterranea CASENT0106207 KJ860239 KJ860051 KJ861939 KJ861326 KJ861510 KJ860668 KJ859863 KJ861133 KJ860901 KJ861743 KJ860472
Tropidomyrmex elianae CASENT0189192 KJ860240 KJ860052 KJ861940 KJ861327 KJ861511 KJ860669 KJ859864 KJ861134 KJ860902 KJ861744 KJ860473
Tyrannomyrmex legatus CASENT0106177 KJ860241 KJ860053 KJ861941 KJ861328 KJ861512 KJ860670 KJ859865 KJ861135 KJ860903 KJ861745 KJ860474
Vitsika suspicax CASENT0068044 KJ860242 KJ860054 KJ861942 KJ861329 KJ861513 KJ860671 KJ859866 KJ861136 KJ860904 KJ861746 KJ860475
Vollenhovia emeryi CASENT0010125 EF012949 EF013077 EF013785 EF013205 EF013657 EF013367 EF013529 GU085795 KJ860905 KJ861747 KJ860476
Vombisidris bilongrudi CASENT0200241 KJ860243 KJ860055 KJ861943 KJ861330 KJ861514 KJ860672 KJ859867 KJ861137 KJ860906 KJ861748 KJ860477
Wasmannia auropunctata CASENT0106114 EF012950 EF013078 EF013786 EF013206 EF013658 EF013368 EF013530 KJ861138 KJ860907 KJ861749 KJ860478
Xenomyrmex floridanus CASENT0106053 EF012951 EF013079 EF013787 EF013207 EF013659 EF013369 EF013531 KJ861139 KJ860908 KJ861750 KJ860479
Acanthoponera minor CASENT0039772 EF012825 EF012953 EF013661 EF013081 EF013533 EF013209 EF013371 KJ861140 KJ860909 KJ861751 KJ860480
Amblyopone australis CASENT0106229 KJ860244 KJ860056 KJ861944 KJ861331 KJ861515 KJ860673 KJ859868 KJ861141 KJ860910 KJ861752 KJ860481
Aneuretus simoni CASENT0007014 EF012833 EF012961 EF013669 EF013089 EF013541 EF013220 EF013382 FJ939840 FJ939894 KJ523645 KJ523562
Ankylomyrma coronacantha CASENT0406734 KJ860245 KJ860057 KJ861945 KJ861332 KJ861516 KJ860674 KJ859869 KJ861142 KJ860911 KJ861753 KJ860482
Apterogyna ZA01 CASENT0106304 KJ860246 KJ860058 KJ861946 KJ861333 KJ861517 KJ860675 KJ859870 KJ861143 KJ860912 KJ861754 KJ860483
Dolichoderus pustulatus CASENT0106164 FJ939760 FJ939792 FJ940028 FJ939824 FJ939995 FJ940060 FJ939963 FJ939859 FJ939913 KJ523685 KJ523602
Lasius californicus CASENT0106045 EF012870 EF012998 EF013706 EF013126 EF013578 EF013268 EF013430 KJ523346 KJ523369 KJ523692 KJ523609
Myrmecia pyriformis CASENT0106088 AY703500 AY703567 AY703634 AY703701 AY703768 EF013292 EF013454 FJ939878 FJ939932 KJ523699 KJ523616
Myrmelachista flavocotea CASENT0106049 EF012889 EF013017 EF013725 EF013145 EF013597 EF013295 EF013457 KJ861144 KJ860913 KJ861755 KJ860484
Nothomyrmecia macrops CASENT0106089 AY703501 AY703568 AY703635 AY703702 AY703769 EF013304 EF013466 FJ939880 FJ939934 KJ523703 KJ523620
Paraponera clavata CASENT0106092 AY703489 AY703556 AY703623 AY703690 AY703757 EF013315 EF013477 KJ523355 KJ523370 KJ523704 KJ523621
Platythyrea mocquerysi CASENT0106094 AY867437 AY867453 AY867422 AY867469 AY867484 EF013322 EF013484 KJ861145 KJ860914 KJ861756 KJ860485
Proceratium stictum CASENT0106095 AY703490 AY703557 AY703624 AY703691 AY703758 EF013335 EF013497 KJ861146 KJ860915 KJ861757 KJ860486
Rhytidoponera chalybaea CASENT0106000 EF012930 EF013058 EF013766 EF013186 EF013638 EF013343 EF013505 FJ939885 FJ939939 KJ523705 KJ523622
Stigmatomma pallipes CASENT0106070 AY703487 AY703554 AY703621 AY703688 AY703755 EF013219 EF013381 KJ523366 KJ523371 KJ523716 KJ523633
Tatuidris tatusia CASENT0423526 EF012939 EF013067 EF013775 EF013195 EF013647 EF013354 EF013516 KJ861147 KJ860916 KJ861758 KJ860487
Tatuidris tatusia_2 CASENT0106203 KJ860247 KJ860059 KJ861947 KJ861334 KJ861518 KJ860676 KJ859871 KJ861148 KJ860917 KJ861759 KJ860488
Tetraponera rufonigra CASENT0106099 AY703515 AY703582 AY703649 AY703716 AY703783 EF013362 EF013524 FJ939892 FJ940011 KJ523718 KJ523635

5
Ward, P.S., Brady, S.G., Fisher, B.L., & Schultz, T.R. (2014) The evolution of myrmicine ants: phylogeny
and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology

Table S2. The 25 "subjective" partitions used in initial Bayesian analyses of the concatenated data,
resulting in a tree that was submitted to PartitionFinder in order to obtain more "objective" partitions.
Models (column 2) were obtained for each partition using JModelTest and, when identified models were
unavailable in MrBayes, the next most complex model was employed (column 3). AA = Abd-A, F2 =
EF1aF2, LR = LW Rh, AK = ArgK, TP = Top1, UB = Ubx, WG = Wg, F1 = EF1aF1, CD = CAD.

Partition Model (from JModelTest) Model Used


AApos1&2 K80+G K80+G
AApos3 GTR+I+G GTR+I+G
F2pos1&2 K80+I+G K80+I+G
F2pos3 TIM2+I+G GTR+I+G
LRpos1 TPM2uf+I+G GTR+I+G
LRpos2 TVM+I+G GTR+I+G
LRpos3 TPM1uf+I+G GTR+I+G
AKpos1 SYM+I+G SYM+I+G
AKpos2 GTR+I+G GTR+I+G
AKpos3 TrN+I+G GTR+I+G
TPpos1 TIM3+I+G GTR+I+G
TPpos2 TVM+I+G GTR+I+G
TPpos3 GTR+I+G GTR+I+G
UBpos1&2 K80+G K80+G
UBpos3 GTR+G GTR+G
WGpos1 GTR+I+G GTR+I+G
WGpos2 GTR+I+G GTR+I+G
WGpos3 TrN+I+G GTR+I+G
F1pos1&2 TrN+I+G GTR+I+G
F1pos3 TPM2uf+G GTR+G
18S TIM1ef+I+G SYM+I+G
28S TIM2+I+G GTR+I+G
CDpos1 TPM3uf+I+G GTR+I+G
CDpos2 HKY+I+G HKY+I+G
CDpos3 TPM2uf+I+G GTR+I+G
Ward, P.S., Brady, S.G., Fisher, B.L., & Schultz, T.R. (2014) The evolution of myrmicine ants: phylogeny
and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology

Table S3a. The "subjective" partitions used in initial, separate Bayesian analyses of each of the 11 genes.
18S and 28S were each analyzed as a single partition. The resulting single-gene trees were submitted to
PartitionFinder in order to obtain more "objective" partitions. Models (column 2) were obtained for
each partition using JModelTest and, when identified models were unavailable in MrBayes, the next
most complex model was employed (column 3). AA = Abd-A, F2 = EF1aF2, LR = LW Rh, AK = ArgK, TP =
Top1, UB = Ubx, WG = Wg, F1 = EF1aF1, CD = CAD.

Subjective Partition JModelTest Model Model Used


AApos1&2 K80+G K80+G
AApos3 GTR+I+G GTR+I+G
AKpos1 SYM+I+G SYM+I+G
AKpos2 GTR+I+G GTR+I+G
AKpos3 TrN+I+G GTR+I+G
CDpos1 TPM3uf+I+G GTR+I+G
CDpos2 HKY+I+G HKY+I+G
CDpos3 TPM2uf+I+G GTR+I+G
F1pos1&2 TrN+I+G GTR+I+G
F1pos3 TPM2uf+G GTR+G
F2pos1&2 K80+I+G K80+I+G
F2pos3 TIM2+I+G GTR+I+G
LRpos1 TPM2uf+I+G GTR+I+G
LRpos2 TVM+I+G GTR+I+G
LRpos3 TPM1uf+I+G GTR+I+G
TPpos1 SYM+I+G SYM+I+G
TPpos2 HKY+I+G HKY+I+G
TPpos3 SYM+I+G SYM+I+G
UBpos1&2 K80+G K80+G
UBpos3 GTR+G GTR+G
WGpos1 GTR+I+G GTR+I+G
WGpos2 GTR+I+G GTR+I+G
WGpos3 TrN+I+G GTR+I+G
18S TIM1ef+I+G SYM+I+G
28S TIM2+I+G GTR+I+G

1
Table S3b. The "objective" partitions and models used in the final, separate Bayesian analyses of each of
the 11 genes. For each of the 9 protein-coding genes, partitions and models were identified by
PartitionFinder ("models = mrbayes") based on an input user tree resulting from a prior Bayesian
analysis employing "subjective" partitions (Table S3a). 18S and 28S were each analyzed as a single
partition with models obtained from JModelTest. AA = Abd-A, F2 = EF1aF2, LR = LW Rh, AK = ArgK, TP =
Top1, UB = Ubx, WG = Wg, F1 = EF1aF1, CD = CAD.

PartitionFinder Partition PartitionFinder Model


AApos1 SYM+I+G
AApos2 F81+I+G
AApos3 GTR+I+G
AKpos1 SYM+I+G
AKpos2 SYM+I+G
AKpos3 SYM+I+G
CDpos1 K80+I+G
CDpos2 GTR+I+G
CDpos3 GTR+I+G
F1pos1 GTR+G
F1pos2 JC
F1pos3 K80+I+G
F2pos1 SYM+I+G
F2pos2 GTR+I+G
F2pos3 SYM+I+G
LRpos1 GTR+I+G
LRpos2 GTR+I+G
LRpos3 SYM+I+G
TPpos1 SYM+I+G
TPpos2 HKY+I+G
TPpos3 SYM+I+G
UBpos1 SYM+I+G
UBpos2 F81+I+G
UBpos3 SYM+G
WGpos1&2 SYM+I+G
WGpos3 SYM+I+G
18S SYM+I+G
28S GTR+I+G

2
Ward, P.S., Brady, S.G., Fisher, B.L., & Schultz, T.R. (2014) The evolution of myrmicine ants: phylogeny
and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology

Table S5. Species distribution matrix, based on the following six biogeographic areas: Neotropical (T),
Nearctic (N), Palearctic (P), Afrotropical (E), Indomalayan (O), and Australasian regions (A). Note that
most species occur in only one region; seven species occupy two regions; one species
(Pristomyrmex_punctatus) inhabits three regions, and none occurs in more than three regions. Tramp
species were assigned to the area in which they are inferred to have originated, based on various
sources including Bolton (1987, 2000), Seifert (2003), and Wilson & Taylor (1967)1.

Taxon T N P E O A
Acanthognathus_ocellatus 1 0 0 0 0 0
Acanthomyrmex_ferox 0 0 0 0 1 0
Acromyrmex_versicolor 1 0 0 0 0 0
Adelomyrmex_paratristani 1 0 0 0 0 0
Adelomyrmex_SC01 0 0 0 1 0 0
Adlerzia_froggatti 0 0 0 0 0 1
Allomerus_octoarticulatus 1 0 0 0 0 0
Ancyridris_polyrhachioides_nr 0 0 0 0 0 1
Anergates_atratulus 0 0 1 0 0 0
Anillomyrma_AFRC_TZ_01 0 0 0 1 0 0
Anillomyrma_decamera 0 0 0 0 1 0
Anisopheidole_antipodum 0 0 0 0 0 1
Aphaenogaster_albisetosa 0 1 0 0 0 0
Aphaenogaster_araneoides 1 0 0 0 0 0
Aphaenogaster_occidentalis 0 1 0 0 0 0
Aphaenogaster_pythia 0 0 0 0 0 1
Aphaenogaster_swammerdami 0 0 0 1 0 0
Apterostigma_auriculatum 1 0 0 0 0 0
Atopomyrmex_mocquerysi 0 0 0 1 0 0
Atta_texana 1 0 0 0 0 0
Austromorium_flavigaster 0 0 0 0 0 1
Baracidris_sitra 0 0 0 1 0 0
Bariamyrma_hispidula 1 0 0 0 0 0
Basiceros_manni 1 0 0 0 0 0
Blepharidatta_conops 1 0 0 0 0 0
Calyptomyrmex_kaurus 0 0 0 1 0 0
Cardiocondyla_mauritanica 0 0 1 0 0 0
Cardiocondyla_MY01 0 0 0 0 1 0
Cardiocondyla_thoracica 0 0 0 0 0 1
Carebara_affinis 0 0 0 0 1 1
Carebara_alperti 0 0 0 0 1 0
Carebara_AU01 0 0 0 0 0 1
Carebara_nosindambo 0 0 0 1 0 0
Carebara_urichi 1 0 0 0 0 0

1
Carebara_vidua 0 0 0 1 0 0
Carebarella_PA01 1 0 0 0 0 0
Cataulacus_ebrardi 0 0 0 1 0 0
Cataulacus_hispidulus 0 0 0 0 1 0
Cephalotes_atratus 1 0 0 0 0 0
Cephalotes_texanus 0 1 0 0 0 0
Chalepoxenus_kutteri 0 0 1 0 0 0
Colobostruma_unicorna 0 0 0 0 0 1
Crematogaster_acuta 1 0 0 0 0 0
Crematogaster_emeryana 0 1 0 0 0 0
Crematogaster_kelleri 0 0 0 1 0 0
Crematogaster_modiglianii 0 0 0 0 1 0
Crematogaster_sordidula 0 0 1 0 0 0
Cryptomyrmex_boltoni 1 0 0 0 0 0
Cyphoidris_exalta 0 0 0 1 0 0
Cyphomyrmex_cornutus 1 0 0 0 0 0
Cyphomyrmex_longiscapus_cf 1 0 0 0 0 0
Dacatria_templaris 0 0 1 0 1 0
Dacetinops_ignotus_cf 0 0 0 0 0 1
Daceton_armigerum 1 0 0 0 0 0
Diaphoromyrma_sofiae 1 0 0 0 0 0
Dicroaspis_KM01 0 0 0 1 0 0
Dilobocondyla_borneensis_cf 0 0 0 0 1 1
Diplomorium_longipenne 0 0 0 1 0 0
Dolopomyrmex_pilatus 0 1 0 0 0 0
Epopostruma_monstrosa 0 0 0 0 0 1
Eurhopalothrix_australis 0 0 0 0 0 1
Eurhopalothrix_gravis 1 0 0 0 0 0
Eutetramorium_mocquerysi 0 0 0 1 0 0
Formicoxenus_diversipilosus 0 1 0 0 0 0
Gauromyrmex_acanthinus 0 0 0 0 1 0
Goniomma_blanci 0 0 1 0 0 0
Harpagoxenus_sublaevis 0 0 1 0 0 0
Huberia_brounii 0 0 0 0 0 1
Hylomyrma_blandiens 1 0 0 0 0 0
Kalathomyrmex_emeryi 1 0 0 0 0 0
Kartidris_sparsipila 0 0 0 0 1 0
Kempfidris_inusualis 1 0 0 0 0 0
Lachnomyrmex_scrobiculatus 1 0 0 0 0 0
Lenomyrmex_colwelli 1 0 0 0 0 0
Leptothorax_muscorum_complex 0 1 1 0 0 0
Liomyrmex_gestroi 0 0 0 0 1 1
Lophomyrmex_ambiguus 0 0 0 0 1 0
Lordomyrma_bhutanensis_cf 0 0 0 0 1 0
Lordomyrma_desupra 0 0 0 0 0 1

2
Machomyrma_dispar 0 0 0 0 0 1
Malagidris_belti 0 0 0 1 0 0
Malagidris_jugum 0 0 0 1 0 0
Manica_bradleyi 0 1 0 0 0 0
Manica_rubida 0 0 1 0 0 0
Mayriella_ebbei 0 0 0 0 0 1
Megalomyrmex_silvestrii 1 0 0 0 0 0
Melissotarsus_insularis 0 0 0 1 0 0
Meranoplus_castaneus 0 0 0 0 1 0
Meranoplus_radamae 0 0 0 1 0 0
Mesostruma_browni 0 0 0 0 0 1
Messor_andrei 0 1 0 0 0 0
Messor_denticornis 0 0 0 1 0 0
Messor_wasmanni 0 0 1 0 0 0
Metapone_madagascarica 0 0 0 1 0 0
Metapone_PG01 0 0 0 0 0 1
Microdaceton_tibialis 0 0 0 1 0 0
Monomorium_antarcticum 0 0 0 0 0 1
Monomorium_brocha 0 0 0 0 1 0
Monomorium_criniceps 0 0 0 0 1 0
Monomorium_denticulatum 1 0 0 0 0 0
Monomorium_destructor 0 0 0 1 0 0
Monomorium_ergatogyna 0 1 0 0 0 0
Monomorium_exiguum 0 0 0 1 0 0
Monomorium_ferodens 0 0 0 1 0 0
Monomorium_fieldi_nr 0 0 0 0 0 1
Monomorium_fisheri 0 0 0 1 0 0
Monomorium_hanneli 0 0 0 1 0 0
Monomorium_latinode 0 0 0 0 1 0
Monomorium_madecassum 0 0 0 1 0 0
Monomorium_MG04 0 0 0 1 0 0
Monomorium_pharaonis 0 0 0 0 1 0
Monomorium_robustior 0 0 0 1 0 0
Monomorium_sechellense 0 0 0 1 0 0
Mycetarotes_acutus 1 0 0 0 0 0
Mycetosoritis_hartmanni 1 1 0 0 0 0
Mycocepurus_goeldii 1 0 0 0 0 0
Myrmecina_americana 0 1 0 0 0 0
Myrmecina_graminicola 0 0 1 0 0 0
Myrmecina_PG01 0 0 0 0 0 1
Myrmica_kotokui 0 0 1 0 0 0
Myrmica_rugiventris 0 1 0 0 0 0
Myrmica_striolagaster 0 1 0 0 0 0
Myrmica_tahoensis 0 1 0 0 0 0
Myrmicaria_carinata 0 0 0 0 1 0

3
Myrmicaria_exigua 0 0 0 1 0 0
Myrmicine_genus26_PH02 0 0 0 0 1 0
Myrmicocrypta_infuscata_cf 1 0 0 0 0 0
Myrmisaraka_brevis 0 0 0 1 0 0
Myrmoxenus_bernardi 0 0 1 0 0 0
Nesomyrmex_echinatinodis 1 0 0 0 0 0
Nesomyrmex_madecassus 0 0 0 1 0 0
Nesomyrmex_MG01 0 0 0 1 0 0
Nesomyrmex_MG08 0 0 0 1 0 0
Nesomyrmex_wilda 1 1 0 0 0 0
Ochetomyrmex_neopolitus 1 0 0 0 0 0
Ochetomyrmex_semipolitus 1 0 0 0 0 0
Octostruma_EC01 1 0 0 0 0 0
Ocymyrmex_fortior_cf 0 0 0 1 0 0
Orectognathus_versicolor 0 0 0 0 0 1
Oxyepoecus_vezenyii 1 0 0 0 0 0
Oxyopomyrmex_santschii 0 0 1 0 0 0
Paratopula_TH01 0 0 0 0 1 0
Perissomyrmex_snyderi 1 0 0 0 0 0
Phalacromyrmex_fugax 1 0 0 0 0 0
Pheidole_clydei 0 1 0 0 0 0
Pheidole_fimbriata 1 0 0 0 0 0
Pheidole_hyatti 0 1 0 0 0 0
Pheidole_longispinosa 0 0 0 1 0 0
Pheidole_lucioccipitalis 0 0 0 0 1 0
Pheidole_pallidula 0 0 1 0 0 0
Pheidole_rhea 0 1 0 0 0 0
Pilotrochus_besmerus 0 0 0 1 0 0
Podomyrma_silvicola 0 0 0 0 0 1
Poecilomyrma_myrmecodiae 0 0 0 0 0 1
Pogonomyrmex_angustus 1 0 0 0 0 0
Pogonomyrmex_imberbiculus 0 1 0 0 0 0
Pogonomyrmex_striatinodus 1 0 0 0 0 0
Pogonomyrmex_subdentatus 0 1 0 0 0 0
Pogonomyrmex_vermiculatus 1 0 0 0 0 0
Pristomyrmex_orbiceps 0 0 0 1 0 0
Pristomyrmex_punctatus 0 0 1 0 1 1
Proatta_butteli 0 0 0 0 1 0
Procryptocerus_scabriusculus 1 0 0 0 0 0
Propodilobus_pingorum 0 0 0 0 1 0
Protalaridris_armata 1 0 0 0 0 0
Recurvidris_TH01 0 0 0 0 1 0
Rhopalomastix_rothneyi 0 0 0 0 0 1
Rhopalothrix_isthmica 1 0 0 0 0 0
Rhoptromyrmex_transversinodis 0 0 0 1 0 0

4
Rogeria_nevadensis 1 0 0 0 0 0
Rogeria_stigmatica 0 0 0 0 0 1
Romblonella_scrobifera 0 0 0 0 0 1
Rotastruma_recava 0 0 0 0 1 0
Rotastruma_stenoceps 0 0 0 0 1 0
Royidris_longiseta 0 0 0 1 0 0
Royidris_notorthotenes 0 0 0 1 0 0
Secostruma_MY01 0 0 0 0 1 0
Solenopsis_fugax 0 0 1 0 0 0
Solenopsis_mameti 0 0 0 1 0 0
Solenopsis_molesta 0 1 0 0 0 0
Solenopsis_papuana 0 0 0 0 0 1
Solenopsis_succinea 1 0 0 0 0 0
Solenopsis_xyloni 0 1 0 0 0 0
Solenopsis_ZA02 0 0 0 1 0 0
Stegomyrmex_manni 1 0 0 0 0 0
Stenamma_dyscheres 0 1 0 0 0 0
Stenamma_expolitum 1 0 0 0 0 0
Stenamma_striatulum 0 0 1 0 0 0
Strongylognathus_testaceus 0 0 1 0 0 0
Strumigenys_ambatrix 0 0 0 1 0 0
Strumigenys_biolleyi 1 0 0 0 0 0
Strumigenys_chiricahua 0 1 0 0 0 0
Strumigenys_chyzeri 0 0 0 0 0 1
Strumigenys_coveri 0 0 0 1 0 0
Strumigenys_dicomas 0 0 0 1 0 0
Strumigenys_emmae 0 0 0 0 0 1
Strumigenys_erynnes 0 0 0 1 0 0
Strumigenys_exiguaevitae 0 0 0 1 0 0
Strumigenys_godeffroyi 0 0 0 0 1 0
Strumigenys_gundlachi 1 0 0 0 0 0
Strumigenys_ludovici 0 0 0 1 0 0
Strumigenys_maxillaris 0 0 0 1 0 0
Strumigenys_membranifera 0 0 0 1 0 0
Strumigenys_nitens 1 0 0 0 0 0
Strumigenys_ocypete 0 0 0 0 1 0
Strumigenys_olsoni 0 0 0 1 0 0
Strumigenys_rogeri 0 0 0 1 0 0
Talaridris_mandibularis 1 0 0 0 0 0
Teleutomyrmex_kutteri 0 0 1 0 0 0
Temnothorax_poeyi 1 0 0 0 0 0
Temnothorax_rugatulus 0 1 0 0 0 0
Terataner_bottegoi 0 0 0 1 0 0
Terataner_MG02 0 0 0 1 0 0
Terataner_MG27 0 0 0 1 0 0

5
Tetheamyrma_subspongia 0 0 0 0 1 0
Tetramorium_caespitum_cf 0 0 1 0 0 0
Tetramorium_CF01 0 0 0 1 0 0
Tetramorium_MG125 0 0 0 1 0 0
Tetramorium_severini 0 0 0 1 0 0
Tetramorium_spinosum 0 1 0 0 0 0
Tetramorium_taylori 0 0 0 0 0 1
Tetramorium_validiusculum 0 0 0 0 0 1
Tetramorium_venator 0 0 0 1 0 0
Trachymyrmex_arizonensis 0 1 0 0 0 0
Tranopelta_gilva 1 0 0 0 0 0
Tranopelta_subterranea 1 0 0 0 0 0
Tropidomyrmex_elianae 1 0 0 0 0 0
Tyrannomyrmex_legatus 0 0 0 0 1 0
Vitsika_suspicax 0 0 0 1 0 0
Vollenhovia_emeryi 0 0 1 0 0 0
Vombisidris_bilongrudi 0 0 0 0 0 1
Wasmannia_auropunctata 1 0 0 0 0 0
Xenomyrmex_floridanus 0 1 0 0 0 0

1
References cited:

Bolton, B. (1987) A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr
(Hymenoptera: Formicidae). Bulletin of the British Museum (Natural History). Entomology, 54, 263–
452.
Bolton, B. (2000) The ant tribe Dacetini. Memoirs of the American Entomological Institute, 65, 1–1028.
Seifert, B. (2003) The ant genus Cardiocondyla (Insecta: Hymenoptera: Formicidae) - a taxonomic
revision of the C. elegans, C. bulgarica, C. batesii, C. nuda, C. shuckardi, C. stambuloffii, C.
wroughtonii, C. emeryi, and C. minutior species groups. Annalen des Naturhistorischen Museums in
Wien. B, Botanik, Zoologie, 104, 203–338.
Wilson, E.O. & Taylor, R.W. (1967) The ants of Polynesia (Hymenoptera: Formicidae). Pacific Insects
Monograph, 14, 1–109.

6
Ward, P.S., Brady, S.G., Fisher, B.L., & Schultz, T.R. (2014) The evolution of myrmicine ants: phylogeny
and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology

Table S6. Support values from the various treatments for selected clades of interest. Treatments
include maximum-likelihood analyses of the full data set (BS, bootstrap proportions); Bayesian analysis
of the full data set (PP, posterior probabilities); (i) "partial RY coding"; (ii) "complete RY coding"; (iii)
exclusion of a minimum subset of 324 characters (not necessarily third positions) deviating significantly
from observed base-frequency averages based on the results of a likelihood-based method (D. Swofford,
unpublished ); and (iv) exclusion of 843 characters (not necessarily third positions), the 10% deviating
the most from observed base-frequency averages, also based on based on the results of a likelihood-
based method for filtering potential nonstationary sites, to be described elsewhere (D. Swofford, pers.
comm.). See text for details. "0+" indicates that a clade is compatible with the most likely/probable
topology but with support 0; "0-" indicates that a clade is incompatible with the most likely/probable
tree, i.e., that it is contradicted by a more likely/probable clade. Taxa that are most unstable across
treatments include Tetheamyrma, Myrmecina, Recurvidris, Adlerzia, Kartidris, Lophomyrmex,
Acanthomyrmex, Mayriella, Nesomyrmex, Xenomyrmex, and Dacetinops.

Clade BS PP PP (i) PP (ii) PP (iii) PP (iv)


Wasmannia + Blepharidatta + Allomerus 98 1.00 1.00 1.00 1.00 1.00
Ochetomyrmex (2 spp) + Tranopelta (2 100 1.00 1.00 1.00 1.00 1.00
spp)
Diaphoromyrma + Lachnomyrmex + 22 0.91 0- 0- 0.53 0-
Wasmannia + Blepharidatta + Allomerus
All Crematogastrini except Tetheamyrma 20 0.90 0- 0- 0- 0.60
Monomorium robustior + M. criniceps + 100 1.00 1.00 1.00 1.00 1.00
M. destructor
Monomorium robustior + M. criniceps + 92 1.00 0.99 0.99 1.00 1.00
M. destructor + Huberia + Podomyrma +
Liomyrmex + Metapone (2 spp) +
Eutetramorium genus-group (7 spp)
Eutetramorium genus-group (7 spp) 92 1.00 0- 0- 1.00 1.00
Cardiocondyla (3 spp) + Ocymyrmex + 0- 0.81 0.69 0.54 0.96 0-
Cataulacus (2 spp) + Atopomyrmex
Huberia + Podomyrma + Liomyrmex + 99 1.00 1.00 1.00 1.00 1.00
Metapone (2 spp)
Perissomymrex + Pristomyrmex (2 spp) 65 1.00 0.94 0.94 1.00 0.99
Melissotarsus + Rhopalomastix + 17 0.99 0- 0- 0- 0-
Terataner (3 spp) + Kartidris + Cyphoidris
+ Dicroaspis + Calyptomyrmex +
Myrmicine genus 26 PH02 + Vollenhovia
+ Tetramorium s.l. (12 spp)
Melissotarsus + Rhopalomastix + 54 1.00 0.61 0.49 0.98 1.00
Terataner (3 spp)
Melissotarsus + Rhopalomastix 96 1.00 1.00 1.00 1.00 1.00

1
Kartidris + Cyphoidris + Dicroaspis + 23 0.97 0- 0- 0- 0-
Calyptomyrmex + Myrmicine genus 26
PH02 + Vollenhovia + Tetramorium s.l.
(12 spp)
Cyphoidris + Dicroaspis + Calyptomyrmex 48 1.00 0.93 0.89 0.97 0-
+ Myrmicine genus 26 PH02 +
Vollenhovia + Tetramorium s.l. (12 spp)
Calyptomyrmex + Myrmicine genus 26 86 1.00 0.36 0- 0.98 0.98
PH02 + Vollenhovia + Tetramorium s.l.
(12 spp)
Calyptomyrmex + Myrmicine genus 26 96 1.00 0.94 0.96 1.00 1.00
PH02 + Vollenhovia
Cyphoidris + Dicroaspis 75 1.00 0- 0- 1.00 0.87
Dacatria + Proatta 75 1.00 1.00 1.00 1.00 1.00
Dilobocondyla + Secostruma 100 1.00 1.00 1.00 1.00 1.00
Lophomyrmex + Adlerzia + Recurvidris 26 0.68 0- 0- 0- 0-
Lophomyrmex + Paratopula + 0- 0- 0.55 0.66 0- 0-
Poecilomyrma + Romblonella +
Rotastruma (2 spp)
Adlerzia + Recurvidris 48 0.91 0- 0- 0.82 0-
Poecilomyrma + Romblonella + 99 1.00 1.00 1.00 1.00 1.00
Rotastruma (2 spp)
Diplomorium + Carebara (6 spp) 99 1.00 1.00 1.00 1.00 1.00
Carebara (6 spp) 99 1.00 1.00 1.00 1.00 1.00
Meranoplus (2 spp) + Crematogaster (5 18 0.93 0- 0- 0.99 0-
spp)
Meranoplus (2 spp) + Recurvidris 0- 0- 0.67 0.70 0- 0-

2
Ward, P.S., Brady, S.G., Fisher, B.L., & Schultz, T.R. (2014) The evolution of myrmicine ants: phylogeny
and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology

Table S7. Range inheritance inferences for major clades of Myrmicinae, based on LAGRANGE analysis.
The range inheritance scenario is expressed as [left|right], where “left” and “right” are the ranges
inherited by the descendant upper branch and the descendant lower branch, respectively, of each clade
depicted in Fig. 3. Only splits (scenarios) with relative probability > 0.05 are given. The ancestral ranges
are abbreviated as follows: T, Neotropical; N, Nearctic; P, Palearctic; E, Afrotropical; O, Indomalayan;
and A, Australasian.

Clade Range relative probability lnL


Myrmicinae T|N 0.204 -558.5
T|NP 0.196 -558.6
T|TNP 0.088 -559.4
TN|N 0.063 -559.7
TE|N 0.061 -559.7
TNE|N 0.052 -559.9
T|TN 0.052 -559.9
Myrmicini N|N 0.338 -558.0
NP|N 0.099 -559.3
N|NP 0.094 -559.3
NP|P 0.089 -559.4
P|P 0.087 -559.4
P|NP 0.084 -559.4
all myrmicines except Myrmicini T|T 0.619 -557.4
TE|T 0.098 -559.3
TN|T 0.074 -559.5
TNE|T 0.050 -559.9
Pogonomyrmecini T|T 0.934 -557.0
Stenammini TNE|N 0.171 -558.7
TNA|N 0.164 -558.7
TE|N 0.157 -558.8
TN|N 0.100 -559.2
E|TN 0.056 -559.8
Solenopsidini T|T 0.973 -557.0
Attini T|T 0.982 -557.0
Crematogastrini TEO|O 0.427 -557.8
EO|O 0.368 -557.9