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Systematic Entomology (2015), 40, 61–81 DOI: 10.1111/syen.12090

The evolution of myrmicine : phylogeny and biogeography of a hyperdiverse (: Formicidae)

PHILIP S. WARD1, SEÁN G. BRADY2, BRIAN L. FISHER3 andTED R. SCHULTZ2 1Department of Entomology and Nematology, University of California, Davis, CA, U.S.A., 2Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A. and 3Department 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 (Hymenoptera: Formicidae), based on analyses of a data matrix comprising 251 and 11 nuclear gene fragments. Under both maximum likelihood and Bayesian methods of inference, we recover a robust phylogeny that reveals six major of Myrmicinae, here treated as newly defined tribes and occur- ring as a pectinate series: , Pogonomyrmecini trib.n., , Solenop- sidini, Attini and . Because we condense the former 25 myrmicine tribes into a new six- scheme, membership in some tribes is now notably different, espe- cially regarding Attini. We demonstrate that the monotypic 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 (). Several species-rich myrmicine genera are shown to be nonmonophyletic, including Pogono- myrmex, , , , , 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. = syn.n. = Protomognathus syn.n.; = Rhoptromyrmex syn.n. = Anergates syn.n. = Teleutomyrmex syn.n. The genus 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; stat.r. is resurrected from synonymy under Monomorium to contain the species in the scabriceps- and destruc- tor-groups; and the monotypic genus 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 or earlier, a number of prominent, species-rich genera, such as Pheidole, , , 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 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: [email protected]

© 2014 The Royal Entomological Society 61 62 P. S . Wa rd 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 (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 of ants (Bolton, 2014; Brady et al., 2014), but four of these – , Formic- selection inae, Myrmicinae and – 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 ( 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 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), (Smith et al., 2011), 1-alpha F1 copy (EF1aF1), elongation factor 1-alpha F2 copy (Nygaard et al., 2012) and (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 sites, 2895 parsimony-informative sites). This matrix (251 taxa p17 EF1aF1 pos3 TIM+G 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 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 . Wa rd 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 ferox, employs the weighted likelihood bootstrap estimator of New- ebbei and floridanus; and (ii) 243 taxa, exclud- ton & Raftery (1994) as modified by Suchard et al. (2001). For ing mauritanica, Cardiocondyla MY01, Cardio- the full concatenated dataset, stationarity was regularly obtained condyla thoracica, gestroi, insularis, within the first few million generations, even for the various madagascarica, Metapone PG01 and 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 aprioriage distributions for 27 Phylogenetic results nodes from the 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 and Rhytidoponera, is the sister ary between the last two. To estimate ancestral areas we group of Myrmicinae, to the exclusion of the . 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 . Wa rd et al.

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

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

Acanthognathus 15 40 73 poinari in Dominican (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 + sculpture to the extant species A. subterranea (Wheeler, 1915). Calibration follows Branstetter (2012). 15 45 73 Two species of Apterostigma in Dominican amber (Schultz, 2007). Median age from Schultz & Brady (2008). 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). 30 45 73 Two species of Cataulacus described from Sicilian amber (Emery, 1891). 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 -group in Dominican amber (Blaimer, 2012b). The calibrated node represents crown Crematogaster (Orthocrema). Median age from Blaimer (2012b). 42 60 85 Myrmica species from Baltic and Saxonian (Dlussky & Rasnitsyn, 2009), not definitely assignable to modern species groups, although similar to the plesiotypic ritae-group (Radchenko et al., 2007). 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. 15 40 73 Undescribed Octostruma in Dominican amber (Wilson, 1988). Pheidole 34 45 73 Pheidole from (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)andHylomyrma. 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 group). 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.Theseare 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). 15 20 73 Trachymyrmex in Dominican amber (Baroni Urbani, 1980). Median age from Schultz & Brady (2008). Outgroups 42 70 90 Stigmatomma groehni in Baltic amber (Dlussky, 2009). The divergence between two subclades of Amblyoponinae (one containing , 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). may be paraphyletic relative to Dolichoderinae, so this fossil calibrates stem dolichoderomorphs. Formicinae 92 95 120 Kyromyrma in New Jersey amber (Grimaldi & Agosti, 2000). 42 60 90 Lasius in Baltic amber (Dlussky, 2011). 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 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 TNPEOA

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 . 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 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 (represented here by Aphaenogaster restricting its membership to these two extant genera, and albisetosa)andVeromessor (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 (, and awaits a more comprehensive investigation of the phylogeny of ) 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 , , Austromorium, , Wardlaw & Elmes, 1998). Similar habits are thought to char- Bariamyrma, Carebarella, , Dolopomyrmex, acterize the genus Manica, although less is known about the , Monomorium, , , biology of these ants (Wheeler & Wheeler, 1970). , Solenopsis, , Tropidomyrmex and Tyran- nomyrmex. The unsampled genus 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 . Wa rd et al.

Strumigenys godeffroyi Strumigenys chyzeri Strumigenys chiricahua 0.96/69 Strumigenys ocypete Strumigenys membranifera Strumigenys exiguaevitae Strumigenys olsoni Strumigenys maxillaris Strumigenys dicomas Strumigenys coveri Carebara affinis Strumigenys rogeri Carebara urichi Strumigenys ludovici Carebara AU01 Strumigenys erynnes Carebara alperti Strumigenys biolleyi Carebara nosindambo Strumigenys gundlachi Strumigenys nitens longipenne Strumigenys ambatrix Cardiocondyla thoracica besmerus Cardiocondyla MY01 fugax 0.96/18 Cardiocondyla mauritanica mandibularis Ocymyrmex fortior cf gravis Cataulacus hispidulus 0.81/44 Eurhopalothrix australis Cataulacus ebrardi Octostruma EC01 mocquerysi manni Nesomyrmex MG08 isthmica Nesomyrmex MG01 armata Nesomyrmex madecassus Pheidole longispinosa Nesomyrmex wilda 0.96/37 Machomyrma dispar Nesomyrmex echinatinodis Anisopheidole antipodum Vitsika suspicax Pheidole lucioccipitalis Myrmisaraka brevis Malagidris belti Pheidole clydei Malagidris jugum Pheidole rhea Royidris longiseta Pheidole fimbriata Royidris notorthotenes 0.99/58 Eutetramorium mocquerysi Cephalotes atratus Metapone PG01 scabriusculus Metapone madagascarica 0.98/16 Trachymyrmex arizonensis Liomyrmex gestroi Atta texana silvicola 0.93/60 Huberia brounii longiscapus cf Monomorium destructor Cyphomyrmex cornutus Monomorium criniceps hartmanni Monomorium robustior acutus ignotus cf emeryi Crematogaster emeryana infuscata cf Crematogaster kelleri goeldii Crematogaster modiglianii Apterostigma auriculatum Crematogaster sordidula 0.93/36 browni 0.93/18 Crematogaster acuta 1.00/59 monstrosa radamae unicorna Meranoplus castaneus 1.00/60 versicolor Pristomyrmex punctatus Pristomyrmex orbiceps 0.95/49 tibialis 1.00/65 colwelli snyderi armigerum Secostruma MY01 0.96/41 borneensis cf auropunctata 0.91/48 TH01 octoarticulatus froggatti conops ambiguus scrobiculatus desupra Diaphoromyrma sofiae Lordomyrma bhutanensis cf gilva Propodilobus pingorum Tranopelta subterranea polyrhachioides nr semipolitus Mayriella ebbei Ochetomyrmex neopolitus Acanthomyrmex ferox Attini Monomorium MG04 graminicola Monomorium exiguum Monomorium fieldi nr Myrmecina PG01 Monomorium ergatogyna Rotastruma stenoceps Monomorium madecassum Rotastruma recava Monomorium pharaonis scrobifera Monomorium hanneli myrmecodiae Cryptomyrmex boltoni TH01 Adelomyrmex SC01 butteli Baracidris sitra templaris Adelomyrmex paratristani Tetramorium spinosum Monomorium fisheri Tetramorium severini Monomorium ferodens Tetramorium MG125 Monomorium sechellense Rhoptromyrmex transversinodis Myrmicaria exigua Teleutomyrmex kutteri Anergates atratulus 0.94/40 Myrmicaria carinata Anillomyrma decamera 0.90/20 testaceus Anillomyrma AFRC TZ 01 cf Tetramorium validiusculum Monomorium brocha Tetramorium venator legatus Tetramorium taylori Tetramorium CF01 emeryi Oxyepoecus vezenyii 1.00/48 Monomorium denticulatum Myrmicine genus26 PH02 0.97/23 kaurus Austromorium flavigaster KM01 Solenopsis succinea exalta 0.99/17 sparsipila Solenopsis xyloni MG27 Solenopsis mameti Terataner MG02 Solenopsis ZA02 1.00/54 Terataner bottegoi 0.99/43 rothneyi Solenopsis papuana Melissotarsus insularis Carebarella PA01 Myrmoxenus bernardi Tropidomyrmex elianae Chalepoxenus kutteri Kempfidris inusualis Rogeria stigmatica Temnothorax poeyi 1.00/67 Bariamyrma hispidula muscorum complex Rogeria nevadensis diversipilosus 0.99/48 Dolopomyrmex pilatus sublaevis Stegomyrmex manni bilongrudi Messor wasmanni acanthinus Solenopsidini Messor denticornis Tetheamyrma subspongia Aphaenogaster occidentalis Crematogastrini Oxyopomyrmex santschii 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 Pogonomyrmex angustus PogonomyrmeciniMyrmica tahoensis Myrmica striolagaster Myrmica rugiventris Myrmicini Manica bradleyi 0.01 length units

0.91/64

Rhytidoponera chalybaea Formicoid Acanthoponera minor flavocotea Lasius californicus macrops pyriformis Tetraponera rufonigra pustulatus Aneuretus simoni Tatuidris tatusia_2 Tatuidris tatusia Poneroid Ankylomyrma coronacantha Platythyrea mocquerysi clavata stictum Stigmatomma pallipes 1.00/62 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-anddestructor-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 aTribal names 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-andsalomonis-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-andsalomonis-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 . Wa rd 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,buttheyalso 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.Thesister 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 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 (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 aprioricalibration 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), (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, (PP91BS48). 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, , 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 ‘’ (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 . 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 . Wa rd et al.

Crematogastrini

Strumigenys chyzeri Strumigenys emmae Strumigenys chiricahua Strumigenys ocypete Strumigenys membranifera Strumigenys exiguaevitae Strumigenys olsoni Strumigenys maxillaris Strumigenys coveri Strumigenys dicomas Strumigenys rogeri Strumigenys erynnes Strumigenys ludovici

Attini 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 Myrmicocrypta infuscata cf Apterostigma auriculatum Epopostruma monstrosa Mesostruma browni Colobostruma unicorna Orectognathus versicolor Microdaceton tibialis 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 Adelomyrmex SC01 Cryptomyrmex boltoni Baracidris sitra Adelomyrmex paratristani Monomorium ferodens Monomorium fisheri sidini Monomorium sechellense

p Anillomyrma decamera Anillomyrma AFRC TZ 01 Myrmicaria carinata Myrmicaria exigua Monomorium brocha Monomorium latinode Tyrannomyrmex legatus

oleno Solenopsis molesta Solenopsis succinea Solenopsidini S 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 Aphaenogaster araneoides StenamminiStenammini Aphaenogaster pythia Aphaenogaster swammerdami Stenamma striatulum Stenamma dyscheres Stenamma expolitum Aphaenogaster albisetosa Messor andrei Pogonomyrmex subdentatus Pogonomyrmex vermiculatus Pogonomyrmex imberbiculus Pogonomyrmecinim Pogonomyrmex striatinodus Hylomyrma blandiens Pogonomyrmex angustus Myrmica striolagaster Myrmica tahoensis MMyrmiciniyrmicini 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 Pristomyrmex orbiceps Crematogastrini 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 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 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 complex other Myrmicinae Formicoxenus diversipilosus 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 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 Taxonomy combinations in Syllophopsis: adiastolon, aureorugosa, aus- 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 . Wa rd et al.

Crematogastrini

Nearctic

Strumigenys chyzeri Neotropical Strumigenys godeffroyi Strumigenys emmae Strumigenys chiricahua Strumigenys ocypete Australasian Strumigenys membranifera Strumigenys exiguaevitae Strumigenys olsoni Indomalayan Strumigenys maxillaris Strumigenys coveri Strumigenys dicomas Afrotropical Strumigenys rogeri Strumigenys erynnes Strumigenys ludovici

Attini Strumigenys gundlachi Palearctic 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 Adelomyrmex SC01 Cryptomyrmex boltoni Baracidris sitra Adelomyrmex paratristani sidini Monomorium ferodens Monomorium fisheri p Monomorium sechellense Anillomyrma decamera Anillomyrma AFRC TZ 01 Myrmicaria carinata Myrmicaria exigua Monomorium brocha Monomorium latinode Tyrannomyrmex legatus oleno Solenopsis molesta Solenopsis succinea S Solenopsidini 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 Aphaenogaster araneoides StenamminiStenammini Aphaenogaster pythia Aphaenogaster swammerdami Stenamma striatulum Stenamma dyscheres Stenamma expolitum Aphaenogaster albisetosa Messor andrei Pogonomyrmex subdentatus Pogonomyrmex vermiculatus PPogonomyrmeciniogonomyrmecini Pogonomyrmex imberbiculus Pogonomyrmex striatinodus Hylomyrma blandiens Pogonomyrmex angustus Myrmica striolagaster Myrmica tahoensis MMyrmiciniyrmicini 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 Nesomyrmex wilda Nearctic Xenomyrmex floridanus Carebara alperti Carebara AU01 Carebara affinis Neotropical Carebara urichi Carebara nosindambo Carebara vidua Australasian Diplomorium longipenne Vitsika suspicax Myrmisaraka brevis Indomalayan Malagidris jugum Malagidris belti Royidris notorthotenes Royidris longiseta Afrotropical Eutetramorium mocquerysi Metapone madagascarica Metapone PG01 Palearctic 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 Propodilobus pingorum Crematogastrini 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 Leptothorax muscorum complex other Myrmicinae Formicoxenus diversipilosus Harpagoxenus sublaevis Vombisidris bilongrudi Fig 3a 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 . Wa rd 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 ( width 0.73–2.90 mm, head length 0.78–2.50 mm); ceps-anddestructor-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; 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 ofa 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 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 ofAphaenogaster ‘ ’ 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, *, 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, *, †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, *, Cyphomyrmex, Daceton, Diaphoromyrma, †Myrmecites, †Orbigastrula, †Quadrulicapito, Epopostruma, Eurhopalothrix,*, Kalathomyrmex, †Quineangulicapito, †Sinomyrmex, †Solenopsites, Lachnomyrmex, Lenomyrmex, Mesostruma, Microdaceton, †Sphaerogasterites, †Wumyrmex and †Zhangidris. *, Mycetarotes,*, Mycetosoritis, Mycocepurus, Myrmicocrypta, Ochetomyrmex, Octostruma, Orectognathus,*Paramycetophylax, Phalacromyrmex, Phei- Excluded from Myrmicinae dole, Pilotrochus, Procryptocerus, Protalaridris,*, Rhopalothrix,*, Strumigenys, Talaridris, Tra- The tribe Ankylomyrmini Bolton and its sole genus, Anky- chymyrmex, Tranopelta and Wasmannia. lomyrma Bolton, are hereby transferred to the subfamily

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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 Figure S1. Majority-rule consensus trees from partitioned, concatenated Bayesian analyses of datasets with ‘wildcard’ References taxon exclusions. Archibald, S.B., Cover, S.P. & Moreau, C.S. (2006) Bulldog ants Figure S2. Majority-rule consensus trees resulting from of the Eocene Okanagan Highlands and history of the subfamily Bayesian analyses of single-gene fragments, under the parti- (Hymenoptera: Formicidae: Myrmeciinae). Annals of the Entomolog- ical Society of America, 99, 487–523. tions and models described in Table S3(b). Aria, C., Perrichot, V. & Nel, A. (2011) Fossil Ponerinae (Hymenoptera: Table S1. List of taxa sampled, voucher specimen codes, and Formicidae) in Early Eocene amber of France. Zootaxa, 2870, 53–62. GenBank accession numbers. Baroni Urbani, C. (1980) First description of fossil gardening ants. (Amber Collection Stuttgart and Natural History Museum Basel; Table S2. The subjective partitions used in initial Bayesian Hymenoptera: Formicidae. I: Attini). Stuttgarter Beiträge zur analyses of the concatenated dataset. Naturkunde. Serie B (Geologie und Paläontologie), 54, 1–13. Baroni Urbani, C. & De Andrade, M.L. (1994) First description of fossil Table S3. The subjective partitions used in initial, separate Dacetini ants with a critical analysis of the current classification of Bayesian analyses of each of the 11 genes (a) and the more the tribe (Amber Collection Stuttgart: Hymenoptera, Formicidae. VI: objective partitions and models used in the final, separate Dacetini). Stuttgarter Beiträge zur Naturkunde. Serie B (Geologie und Bayesian analyses of each of these genes (b). Paläontologie), 198, 1–65. Baroni Urbani, C. & De Andrade, M.L. (2007) The ant tribe Dacetini: Table S4. Summary of rate heterogeneity values for limits and constituent genera, with descriptions of new species third-position partitions of protein-coding genes, which (Hymenoptera, Formicidae). Annali del Museo Civico di Storia informed the ‘partial RY coding’ and ‘complete RY Naturale “Giacomo Doria”, 99, 1–191. Beibl, J., Stuart, R.J., Heinze, J. & Foitzik, S. (2005) Six origins of coding’ analyses. AA = Abd-A,F2= EF1aF2,LR= LW slavery in formicoxenine ants. Insectes Sociaux, 52, 291–297. Rh,AK= ArgK,TP= Top1,UB= Ubx,WG= Wg, Blaimer, B.B. (2012a) Untangling complex morphological variation: F1 = EF1aF1,CD= CAD. taxonomic revision of the subgenus Crematogaster (Oxygyne)in

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81 Phylogeny and evolution of myrmicine ants 79

Madagascar, with insight into the evolution and biogeography of Davidson, D.W. & McKey, D. (1993) Ant-plant symbioses: stalking the this enigmatic ant clade (Hymenoptera: Formicidae). Systematic Chuyachaqui. Trends in Ecology & Evolution, 8, 326–332. Entomology, 37, 240–260. De Andrade, M.L. (1992) First fossil “true Macromischa” in amber from Blaimer, B.B. (2012b) Acrobat ants go global – origin, evolution and the Dominican Republic (Hymenoptera, Formicidae). Mitteilungen systematics of the genus Crematogaster (Hymenoptera: Formicidae). der Schweizerischen Entomologischen Gesellschaft, 65, 341–351. Molecular and Evolution, 65, 421–436. De Andrade, M.L. & Baroni Urbani, C. (1999) Diversity and adaptation Bolton, B. (1973) A remarkable new arboreal ant genus (Hym. Formi- in the ant genus Cephalotes, past and present. Stuttgarter Beiträge zur cidae) from West Africa. Entomologist’s Monthly Magazine, 108, Naturkunde. Serie B (Geologie und Paläontologie), 271, 1–889. 234–237. De Andrade, M.L., Baroni Urbani, C., Brandão, C.R.F. & Wagensberg, Bolton, B. (1987) A review of the Solenopsis genus-group and revision J. (1999) Two new species of Leptothorax “Nesomyrmex” fossils in of Afrotropical Monomorium Mayr (Hymenoptera: Formicidae). Dominican amber (Hymenoptera: Formicidae). Beiträge zur Ento- Bulletin of the British Museum (Natural History). Entomology, 54, mologie, 49, 133–140. 263–452. Dlussky, G.M. (1996) Ants (Hymenoptera: Formicidae) of Burmese Bolton, B. (1994) Identification Guide to the Ant Genera of the World. amber. Paleontologicheskii Zhurnal, 1996, 83–89 (in Russian). Harvard University Press, Cambridge, MA. Dlussky, G.M. (2009) The ant subfamilies Ponerinae, Cerapachyi- Bolton, B. (1999) Ant genera of the tribe Dacetonini (Hymenoptera: nae, and (Hymenoptera, Formicidae) in the Late Formicidae). Journal of Natural History, 33, 1639–1689. Eocene Ambers of Europe. Paleontological Journal, 43, 1043–1086. Bolton, B. (2003) Synopsis and classification of Formicidae. Memoirs Dlussky, G.M. (2011) The ants of the genus Lasius Fabricius of the American Entomological Institute, 71, 1–370. (Hymenoptera, Formicidae) from the Late Eocene European ambers. Bolton, B. (2014) AntCat. An Online Catalog of the Ants of the World Vestnik Zoologii, 45, 209–222. [WWW document]. URL http://antcat.org/ [accessed on 8 May 2014]. Dlussky, G.M. (2012) New fossil ants of the subfamily Myrmeciinae Bolton, B. & Fisher, B.L. (2014) The Madagascan endemic myrmicine (Hymenoptera: Formicidae) from Germany. Paleontological Journal, ants related to Eutetramorium (Hymenoptera: Formicidae): taxonomy 46, 288–292. of the genera Eutetramorium Emery, Malagidris nom. n., Myrmis- Dlussky, G.M. & Radchenko, A.G. (2009) Two new primitive ant araka gen. n., Royidris gen. n., and Vitsika gen. n. Zootaxa, 3791, genera from the late Eocene European ambers. Acta Palaeontologica 1–99. Polonica, 54, 435–441. Brady, S.G., Fisher, B.L., Schultz, T.R. & Ward, P.S. (2014) The rise Dlussky, G.M. & Radchenko, A.G. (2011) Pristomyrmex rasnitsyni sp. of the army ants: diversification of the specialized predatory doryline n., the first known fossil species of the ant genus Pristomyrmex Mayr ants. BMC Evolutionary Biology, 14, 93. (Hymenoptera: Formicidae) from the Late Eocene Danish amber. Brady, S.G., Schultz, T.R., Fisher, B.L. & Ward, P.S. (2006) Evaluating Russian Entomological Journal, 20, 251–254. alternative hypotheses for the early evolution and diversification of Dlussky, G.M. & Rasnitsyn, A.P. (2009) Ants (Insecta: Vespida: Formi- ants. Proceedings of the National Academy of Sciences of the United cidae) in the Upper Eocene amber of central and eastern Europe. States of America, 103, 18 172–18 177. Paleontological Journal, 43, 1024–1042. Branstetter, M.G. (2009) The ant genus Stenamma Westwood Dlussky, G.M., Brothers, D.J. & Rasnitsyn, A.P. (2004) The first Late (Hymenoptera: Formicidae) redefined, with a description of a Cretaceous ants (Hymenoptera: Formicidae) from southern Africa, new genus Propodilobus. Zootaxa, 2221, 41–57. with comments on the origin of the Myrmicinae. Systematics Branstetter, M.G. (2012) Origin and diversification of the cryptic ant and Evolution, 35, 1–13. genus Stenamma Westwood (Hymenoptera: Formicidae), inferred Drummond, A.J., Ho, S.Y.W., Phillips, M.J. & Rambaut, A. (2006) from multilocus molecular data, biogeography and natural history. Relaxed phylogenetics and dating with confidence. PLoS Biology, 4, Systematic Entomology, 37, 478–496. e88. Brian, M.V. (1983) Social Insects. Ecology and Behavioural Biology. Drummond, A.J., Suchard, M.A., Xie, D. & Rambaut, A. (2012) Chapman & Hall, London. Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecular Brown, W.L. Jr. (1986) Indomyrma dasypyx, new genus and species, Biology and Evolution, 29, 1969–1973. a myrmicine ant from peninsular India (Hymenoptera: Formicidae). Dubovikoff, D.A. (2011) The first record of the genus Pheidole West- Israel Journal of Entomology, 19, 37–49. wood, 1839 (Hymenoptera, Formicidae) from the Baltic amber. Rus- Brown, W.L. Jr. (2000) Diversity of ants. Ants. Standard Methods for sian Entomological Journal, 20, 255–257. Measuring and Monitoring (ed. by D. Agosti, J.D. Majer, Elmes, G.W. (1991) The social biology of Myrmica ants. Actes des L.E. Alonso and T.R. Schultz), pp. 45–79. Smithsonian Institution Colloques Insectes Sociaux, 7, 17–34. Press, Washington, DC. Emery, C. (1891) Le formiche dell’ambra Siciliana nel Museo Miner- Brown, J.M., Hedtke, S.M., Lemmon, A.R. & Lemmon, E.M. (2010) alogico dell’Università di Bologna. Memorie della Reale Accademia When trees grow too long: investigating the causes of highly inac- delle Scienze dell’Istituto di Bologna, 5, 141–165. curate Bayesian branch-length estimates. Systematic Biology, 59, Ettershank, G. (1966) A generic revision of the world Myrmicinae 145–161. related to Solenopsis and Pheidologeton (Hymenoptera: Formicidae). Buschinger, A. (2009) Social parasitism among ants: a review Australian Journal of Zoology, 14, 73–171. (Hymenoptera: Formicidae). Myrmecological News, 12, 219–235. Fernández, F. & Palacio, E.E. (1999) Lenomyrmex, an enigmatic new Carpenter, F.M. (1930) The fossil ants of North America. Bulletin of the ant genus from the Neotropical region (Hymenoptera: Formicidae: Museum of Comparative Zoology, 70, 1–66. Myrmicinae). Systematic Entomology, 24, 7–16. Cole, A.C. Jr. (1968) Pogonomyrmex Harvester Ants. A Study of the Fiedler, K. (2006) Ant-associates of Palaearctic lycaenid butterfly larvae Genus in North America. Knoxville, TN, University of Tennessee (Hymenoptera: Formicidae; Lepidoptera: Lycaenidae) – a review. Press. Myrmecologische Nachrichten, 9, 77–87. Cox, C.B. (2001) The biogeographic regions reconsidered. Journal of Fischer, G., Azorsa, F. & Fisher, B.L. (2014) The ant genus Carebara Biogeography, 28, 511–523. Westwood (Hymenoptera: Formicidae): synonymisation of Phei- Davidson, D.W. (1977) Foraging ecology and community organization dologeton Mayr under Carebara, establishment and revision of the in desert seed-eating ants. Ecology, 58, 725–737. C. polita species group. ZooKeys, in press.

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81 80 P. S . Wa rd et al.

Grimaldi, D. & Agosti, D. (2000) A formicine in New Jersey Cretaceous Moreau, C.S. (2008) Unraveling the evolutionary history of the hyper- amber (Hymenoptera: Formicidae) and early evolution of the ants. diverse ant genus Pheidole (Hymenoptera: Formicidae). Molecular Proceedings of the National Academy of Sciences of the United States Phylogenetics and Evolution, 48, 224–239. of America, 97, 13 678–13 683. Moreau, C.S. & Bell, C.D. (2013) Testing the museum versus cradle Heterick, B.E. (2001) Revision of the Australian ants of the genus tropical biological diversity hypothesis: phylogeny, diversification, Monomorium (Hymenoptera: Formicidae). Invertebrate Taxonomy, and ancestral biogeographic range evolution of the ants. Evolution, 15, 353–459. 67, 2240–2257. Heterick, B. (2006) A revision of the Malagasy ants belonging to genus Moreau, C.S., Bell, C.D., Vila, R., Archibald, S.B. & Pierce, N.E. (2006) Monomorium Mayr, 1855 (Hymenoptera: Formicidae). Proceedings Phylogeny of the ants: diversification in the age of angiosperms. of the California Academy of Sciences, 57, 69–202. Science, 312, 101–104. Hölldobler, B. & Wilson, E.O. (1990) The Ants. Harvard University Mueller, U.G., Schultz, T.R., Currie, C.R., Adams, R.M.M. & Malloch, Press, Cambridge, MA. D. (2001) The origin of the attine ant-fungus mutualism. Quarterly Hölldobler, B. & Wilson, E.O. (2011) The Leafcutter Ants. Civilization Review of Biology, 76, 171–197. by Instinct. W. W. Norton, New York, NY. Newton, M.A. & Raftery, A.E. (1994) Approximate Bayesian inference Hong, Y.-C. (2002) Amber Insects of China. Beijing Science and by the weighted likelihood bootstrap. Journal of the Royal Statistical Technology Press, Beijing (in Chinese). Society, Series B, 56, 3–48. Jansen, G. & Savolainen, R. (2010) Molecular phylogeny of the ant Nixon, K.C. & Wheeler, Q.D. (1992) Extinction and the origin of tribe Myrmicini (Hymenoptera: Formicidae). Zoological Journal of species. Extinction and Phylogeny (ed. by Q.D. Wheeler and M. the Linnean Society, 160, 482–495. Novacek), pp. 119–143. Columbia University Press, New York, NY. Jansen, G., Savolainen, R. & Vepsäläinen, K. (2010) Phylogeny, Nygaard, S., Zhang, G., Schiøtt, M. et al. (2012) The genome of the divergence-time estimation, biogeography and social parasite–host leaf-cutting ant suggests key adaptations to relationships of the Holarctic ant genus Myrmica (Hymenoptera: advanced social life and fungus farming. Genome Research, 21, Formicidae). and Evolution, 56, 294–304. 1339–1348. Johnson, R.A. (2001) Biogeography and community structure of North Nylander, J.A.A., Ronquist, F., Huelsenbeck, J.P. & Nieves-Audrey, J.L. American seed-harvesting ants. Annual Review of Entomology, 46, (2004) Bayesian phylogenetic analysis of combined data. Systematic 1–29. Biology, 53, 47–67. Kearney, M. (2002) Fragmentary taxa, missing data, and ambigu- Ogata, K. & Okido, H. (2007) Revision of the ant genus Perissomyrmex ity: mistaken assumptions and conclusions. Systematic Biology, 51, with notes on the phylogeny of the tribe Myrmecinini. Memoirs of the 369–381. American Entomological Institute, 80, 352–369. Keller, R.A. (2011) A phylogenetic analysis of ant morphology Pacheco, J.A. & Mackay, W.P. (2013) The Systematics and Biology of (Hymenoptera: Formicidae) with special reference to the ponero- the New World Thief Ants of the Genus Solenopsis (Hymenoptera: morph subfamilies. Bulletin of the American Museum of Natural Formicidae). Edwin Mellen Press, Lewiston, NY. History, 355, 1–90. Posada, D. (2008) jModelTest: phylogenetic model averaging. Molecu- Kempf, W.W. (1973) A revision of the Neotropical myrmicine ant genus lar Biology and Evolution, 25, 1253–1256. Hylomyrma Forel (Hymenoptera: Formicidae). Studia Entomologica, Posada, D. & Buckley, T.R. (2004) Model selection and model averaging 16, 225–260. in phylogenetics: advantages of the AIC and Bayesian approaches Kugler, C. (1979) Evolution of the sting apparatus in the myrmicine ants. over likelihood ratio tests. Systematic Biology, 53, 793–808. Evolution, 33, 117–130. Price, S.L., Powell, S., Kronauer, D.J.C., Tran, L.A.P., Pierce, N.E. Lach, L., Parr, C.L. & Abbott, K. (eds) (2009) Ant Ecology. Oxford & Wayne, R.K. (2014) Renewed diversification is associated with University Press, Oxford. new ecological opportunity in the Neotropical turtle ants. Journal of Lanfear, R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) Partition- Evolutionary Biology, 27, 242–258. Finder: combined selection of partitioning schemes and substitution Rabeling, C., Brown, J.M. & Verhaagh, M. (2008) Newly discovered models for phylogenetic analyses. Molecular Biology and Evolution, sister lineage sheds light on early ant evolution. Proceedings of the 29, 1695–1701. National Academy of Sciences of the United States of America, 105, LaPolla, J.S., Dlussky, G.M. & Perrichot, V. (2013) Ants and the fossil 14 913–14 917. record. Annual Review of Entomology, 58, 609–630. Radchenko, A.G. (1997) Review of the ants of scabriceps group of Lucky, A. & Sarnat, E.M. (2010) Biogeography and diversification of the genus Monomorium Mayr (Hymenoptera, Formicidae). Annales the Pacific ant genus Lordomyrma Emery. Journal of Biogeography, Zoologici (Warsaw), 46, 211–224. 37, 624–634. Radchenko, A.G., Dlussky, G. & Elmes, G.W. (2007) The ants of the Maddison, D.R. & Maddison, W.P. (2005) MacClade 4: Analysis of Phy- genus Myrmica (Hymenoptera, Formicidae) from Baltic and Saxo- logeny and Character Evolution. Version 4.08 [WWW document]. nian amber (Late Eocene). Journal of Paleontology, 81, 1494–1501. URL http://macclade.org [accessed on 24 March 2014]. Rambaut, A. & Drummond A.J. (2009) Tracer v1.5 [WWW document]. Maddison, W.P. & Maddison, D.R. (2010) Mesquite: A Modular System URL http://beast.bio.ed.ac.uk/Tracer [accessed on 14 January 2014]. for Evolutionary Analysis. Version 2.73. [WWW document]. URL Ree, R.H. & Smith, S.A. (2008) Maximum likelihood inference of geo- http://mesquiteproject.org [accessed on 24 March 2014]. graphic range evolution by dispersal, local extinction, and cladogen- Marshall, D.C. (2010) Cryptic failure of partitioned Bayesian phyloge- esis. Systematic Biology, 57, 4–14. netic analyses: lost in the land of long trees. Systematic Biology, 59, Ree, R.H., Moore, B.R., Webb, C.O. & Donoghue, M.J. (2005) A 108–117. likelihood framework for inferring the evolution of geographic range Marshall, D.C., Simon, C. & Buckley, T.R. (2006) Accurate branch on phylogenetic trees. Evolution, 59, 2299–2311. length estimation in partitioned Bayesian analyses requires accom- Ronquist, F., Teslenko, M., van der Mark, P. et al. (2012) MrBayes 3.2: modation of among-partition rate variation and attention to branch efficient Bayesian phylogenetic inference and model choice across a length priors. Systematic Biology, 55, 993–1003. large model space. Systematic Biology, 61, 539–542. Mayr, G. (1868) Die Ameisen des baltischen Bernsteins. Beiträge zur Russell, J.A., Moreau, C., Goldman-Huertas, B., Fujiwara, M., Lohman, Naturkunde Preussens, 1, 1–102. D. & Pierce, N.E. (2009) Bacterial gut symbionts are tightly

© 2014 The Royal Entomological Society, Systematic Entomology, 40, 61–81 Phylogeny and evolution of myrmicine ants 81

linked with the evolution of herbivory in ants. Proceedings of the Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, National Academy of Sciences of the United States of America, 106, D.G. (1997) The ClustalX Windows interface: flexible strategies for 21 236–21 241. multiple sequence alignment aided by quality analysis tools. Nucleic Rust, J., Singh, H., Rana, R.S. et al. (2010) Biogeographic and Acids Research, 24, 4876–4882. evolutionary implications of a diverse paleobiota in amber Tschinkel, W.R. (2006) The Fire Ants. Harvard University Press, from the early Eocene of India. Proceedings of the National Cambridge, MA. Academy of Sciences of the United States of America, 107, Ward, P.S. (2007a) The ant genus Leptanilloides: discovery of the male 18 360–18 365. and evaluation of phylogenetic relationships based on DNA sequence Schlick-Steiner, B.C., Steiner, F.M., Moder, K., Seifert, B., Sanetra, M., data. Memoirs of the American Entomological Institute, 80, 637–649. Dyreson, E., Stauffer, C. & Christian, E. (2006) A multidisciplinary Ward, P.S. (2007b) Phylogeny, classification, and species-level taxon- approach reveals cryptic diversity in western Palearctic Tetramorium omy of ants (Hymenoptera: Formicidae). Zootaxa, 1668, 549–563. ants (Hymenoptera: Formicidae). Molecular Phylogenetics and Evo- Ward, P.S. (2011) Integrating molecular phylogenetic results into ant lution, 40, 259–273. taxonomy (Hymenoptera: Formicidae). Myrmecological News, 15, Schmidt, C. (2013) Molecular phylogenetics of ponerine ants 21–29. (Hymenoptera: Formicidae: Ponerinae). Zootaxa, 3647, Ward, P.S. & Downie, D.A. (2005) The ant subfamily Pseudomyrmeci- 201–250. nae (Hymenoptera: Formicidae): phylogeny and evolution of big-eyed Schultz, T.R. (2007) The fungus-growing ant genus Apterostigma in arboreal ants. Systematic Entomology, 30, 310–335. Dominican amber. Memoirs of the American Entomological Institute, Ward, P.S. & Sumnicht, T.P. (2012) Molecular and morphological 80, 425–436. evidence for three sympatric species of (Hymenoptera: Schultz, T.R. & Brady, S.G. (2008) Major evolutionary transitions in ant Formicidae) on the Greek island of Rhodes. Myrmecological News, agriculture. Proceedings of the National Academy of Sciences of the 17, 5–11. United States of America, 105, 5435–5440. Ward, P.S., Brady, S.G., Fisher, B.L. & Schultz, T.R. (2010) Phylogeny Scotese, C.R. (2010) PALEOMAP Project [WWW document]. URL and biogeography of dolichoderine ants: effects of data partition- http://www.scotese.com/ [accessed on 24 March 2014]. ing and relict taxa on historical inference. Systematic Biology, 59, Seifert, B. (1988) A taxonomic revision of the Myrmica species 342–362. of Europe, Asia Minor, and Caucasia (Hymenoptera, Formicidae). Wardlaw, J.C. & Elmes, G.W. (1998) Variability in oviposition by Abhandlungen und Berichte des Naturkundemuseums Görlitz, 62, workers of six species of Myrmica (Hymenoptera, Formicidae). 1–75. Insectes Sociaux, 45, 369–384. Seifert, B. (2003) The ant genus Cardiocondyla (Insecta: Hymenoptera: Wallace, A.R. (1876) The Geographical Distribution of . Formicidae) – a taxonomic revision of the C. elegans, C. bulgarica, Macmillan, London. C. batesii, C. nuda, C. shuckardi, C. stambuloffii, C. wroughtonii, Wheeler, G.C. & Wheeler, J. (1970) The natural history of Manica C. emeryi,andC. minutior species groups. Annalen des Naturhis- (Hymenoptera: Formicidae). Journal of the Kansas Entomological torischen Museums in Wien. Serie B, Botanik und Zoologie, 104, Society, 43, 129–162. 203–338. Wheeler, W.M. (1915) The ants of the Baltic Amber. Schriften der Smith, C.R., Smith, C.D., Robertson, H.M. et al. (2011) Draft genome Physikalisch-Ökonomischen Gesellschaft zu Königsberg, 55, 1–142. of the red Pogonomyrmex barbatus. Proceedings of the Wilson, E.O. (1988) The biogeography of the West Indian ants National Academy of Sciences of the United States of America, 108, (Hymenoptera: Formicidae) Zoogeography of Caribbean Insects (ed. 5667–5672. by J.K. Liebherr), pp. 214-230. Cornell University Press, Ithaca, NY. Suchard, M.A., Weiss, R.E. & Sinsheimer, J.S. (2001) Bayesian selec- Wilson, E.O. (1989) Chimaeridris, a new genus of hook-mandibled myr- tion of continuous-time Markov chain evolutionary models. Molecu- micine ants from tropical Asia (Hymenoptera: Formicidae). Insectes lar Biology and Evolution, 18, 1001–1013. Sociaux, 36, 62–69. Suen, G., Teiling, C., Li, L. et al. (2011) The genome sequence of Wilson, E.O. & Hölldobler, B. (2005) The rise of the ants: a phylogenetic the leaf-cutter ant reveals insights into its obligate and ecological explanation. Proceedings of the National Academy of symbiotic lifestyle. PLoS Genetics, 7, e1002007. Sciences of the United States of America, 102, 7411–7414. Swofford, D.S. (2002) PAUP*. Phylogenetic Analysis Using Parsimony Wurm, Y., Wang, J., Riba-Grognuz, O. et al. (2011) The genome of the (*and Other Methods) v. 4.0b10. Sinauer Associates, Sunderland, fire ant Solenopsis invicta. Proceedings of the National Academy of MA. Sciences of the United States of America, 108, 5679–5684. Taber, S.W. (1998) The World of Harvester Ants. Texas A&M University Zwickl, D.J. (2006) Genetic algorithm approaches for the phylogenetic Press, College Station, TX. analysis of large biological sequence datasets under the maximum Taylor, R.W. (2009) Ants of the genus Lordomyrma Emery (1) Generic likelihood criterion. PhD dissertation, The University of Texas at synonymy, composition and distribution, with notes on Ancyridris Austin, Austin, Texas. Wheeler and Cyphoidris Weber (Hymenoptera: Formicidae: Myrmic- inae). Zootaxa, 1979, 16–28. Accepted 9 May 2014 Terayama, M. (2009) A synopsis of the Formicidae of Taiwan First published online 23 July 2014 (Insecta: Hymenoptera). Research Bulletin of Kanto Gakuen Univer- sity. Liberal Arts, 17, 81–266. Terayama, M. & Yamane, S. (2000) Lasiomyrma, a new stenammine ant genus from southeast Asia (Hymenoptera: Formicidae). Entomologi- cal Science, 3, 523–527.

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

Figure S1a 248 taxa

Apterogyna ZA01 100 Amblyopone australis 89 Stigmatomma pallipes 88 Proceratium stictum 58 69 Platythyrea mocquerysi 100 100 Ankylomyrma coronacantha Tatuidris tatusia Tatuidris tatusia_2 100 Aneuretus simoni 100 Dolichoderus pustulatus 100 Tetraponera rufonigra 100 Myrmecia pyriformis Nothomyrmecia macrops 100 100 Lasius californicus Myrmelachista flavocotea 100 Acanthoponera minor Rhytidoponera chalybaea 100 100 Manica bradleyi 100 Manica rubida 100 Myrmica kotokui 100 Myrmica rugiventris 100 Myrmica striolagaster 90 Myrmica tahoensis 100 Pogonomyrmex angustus 99 Hylomyrma blandiens 100 Pogonomyrmex striatinodus 100 Pogonomyrmex imberbiculus 100 Pogonomyrmex subdentatus 100 Pogonomyrmex vermiculatus 100 Aphaenogaster albisetosa 100 Messor andrei 100 100 Stenamma expolitum 100 Stenamma dyscheres Stenamma striatulum 100 Aphaenogaster pythia 100 Aphaenogaster swammerdami 75 Aphaenogaster araneoides 100 Goniomma blanci 100 Oxyopomyrmex santschii 100 Aphaenogaster occidentalis 100 Messor denticornis 97 Messor wasmanni Stegomyrmex manni 99 Dolopomyrmex pilatus 100 Rogeria nevadensis 100 Bariamyrma hispidula 100 Rogeria stigmatica 100 Austromorium flavigaster 85 Monomorium antarcticum 100 Monomorium denticulatum 99 100 Oxyepoecus vezenyii Kempfidris inusualis 100 Tropidomyrmex elianae 100 Carebarella PA01 100 100 Solenopsis papuana 100 Solenopsis fugax 100 Solenopsis ZA02 100 Solenopsis mameti 100 Solenopsis xyloni 100 100 Solenopsis molesta Solenopsis succinea Megalomyrmex silvestrii 100 Tyrannomyrmex legatus 72 79 Monomorium brocha Monomorium latinode 72 Monomorium sechellense 100 100 Monomorium ferodens 87 Monomorium fisheri 100 Adelomyrmex paratristani 100 Baracidris sitra 100 Adelomyrmex SC01 Cryptomyrmex boltoni 100 Anillomyrma AFRC TZ 01 94 Anillomyrma decamera 100 Myrmicaria carinata Myrmicaria exigua 89 Monomorium hanneli 100 Monomorium pharaonis 55 Monomorium madecassum 100 Monomorium ergatogyna 100 Monomorium fieldi nr 100 Monomorium exiguum 100 Monomorium MG04 100 Ochetomyrmex neopolitus 100 Ochetomyrmex semipolitus Tranopelta subterranea 100 Tranopelta gilva 92 Diaphoromyrma sofiae 89 Lachnomyrmex scrobiculatus 100 Blepharidatta conops 100 Allomerus octoarticulatus 100 Wasmannia auropunctata 95 Acanthognathus ocellatus 72 Daceton armigerum 95 Lenomyrmex colwelli 100 Microdaceton tibialis 100 Orectognathus versicolor 100 100 Colobostruma unicorna 94 100 Epopostruma monstrosa Mesostruma browni 100 Apterostigma auriculatum 100 Mycocepurus goeldii 100 Myrmicocrypta infuscata cf 100 Kalathomyrmex emeryi 59 Mycetarotes acutus 100 Mycetosoritis hartmanni 100 Cyphomyrmex cornutus 93 Cyphomyrmex longiscapus cf 100 Acromyrmex versicolor 98 Atta texana Trachymyrmex arizonensis 100 100 Procryptocerus scabriusculus 99 Cephalotes atratus Cephalotes texanus 100 Pheidole fimbriata 100 Pheidole rhea 100 Pheidole clydei 69 Pheidole hyatti 100 Pheidole lucioccipitalis 100 Pheidole pallidula 96 100 Anisopheidole antipodum 52 Machomyrma dispar Pheidole longispinosa 100 Protalaridris armata 100 Rhopalothrix isthmica 90 Basiceros manni 100 Octostruma EC01 100 Eurhopalothrix australis 82 100 100 Eurhopalothrix gravis Talaridris mandibularis 100 Phalacromyrmex fugax 100 Pilotrochus besmerus Strumigenys ambatrix 100 100 100 Strumigenys nitens Strumigenys gundlachi 100 100 Strumigenys biolleyi Strumigenys erynnes 100 Strumigenys ludovici 100 100 Strumigenys rogeri 87 Strumigenys coveri Strumigenys dicomas 100 Strumigenys maxillaris 95 Strumigenys olsoni 96 97 Strumigenys exiguaevitae Strumigenys membranifera 100 Strumigenys ocypete 91 Strumigenys chiricahua 100 Strumigenys emmae 100 Strumigenys chyzeri Strumigenys godeffroyi Tetheamyrma subspongia 100 Gauromyrmex acanthinus Vombisidris bilongrudi 69 100 99 Harpagoxenus sublaevis 74 Formicoxenus diversipilosus Leptothorax muscorum complex 100 Temnothorax poeyi 89 75 Temnothorax rugatulus 100 Chalepoxenus kutteri Myrmoxenus bernardi 100 Melissotarsus insularis 100 Rhopalomastix rothneyi 100 100 Terataner bottegoi 100 Terataner MG02 Terataner MG27 Kartidris sparsipila 100 98 100 Cyphoidris exalta Dicroaspis KM01 100 100 Calyptomyrmex kaurus 100 Myrmicine genus26 PH02 100 Vollenhovia emeryi 100 Tetramorium venator Tetramorium validiusculum 100 Tetramorium CF01 Tetramorium taylori 100 100 Tetramorium caespitum cf 100 Strongylognathus testaceus 100 Anergates atratulus Teleutomyrmex kutteri 100 Rhoptromyrmex transversinodis 97 Tetramorium MG125 100 Tetramorium severini Tetramorium spinosum 87 Dacetinops ignotus cf Lophomyrmex ambiguus 71 Adlerzia froggatti Recurvidris TH01 100 Dacatria templaris Proatta butteli 100 Dilobocondyla borneensis cf Secostruma MY01 100 Myrmecina PG01 100 Myrmecina americana Myrmecina graminicola 100 Perissomyrmex snyderi 100 Pristomyrmex orbiceps Pristomyrmex punctatus 100 Propodilobus pingorum 54 Ancyridris polyrhachioides nr 94 Lordomyrma bhutanensis cf Lordomyrma desupra 100 Paratopula TH01 100 Poecilomyrma myrmecodiae Romblonella scrobifera 100 Rotastruma recava Rotastruma stenoceps 100 Diplomorium longipenne 100 Carebara vidua 100 Carebara nosindambo 66 100 Carebara alperti 100 Carebara AU01 100 Carebara urichi Carebara affinis 100 Meranoplus castaneus 83 Meranoplus radamae 100 Crematogaster acuta 100 Crematogaster sordidula 100 Crematogaster modiglianii 100 Crematogaster adrepens Crematogaster emeryana 100 Nesomyrmex echinatinodis 100 Nesomyrmex wilda 100 Nesomyrmex madecassus 90 94 Nesomyrmex MG01 Nesomyrmex MG08 Atopomyrmex mocquerysi 75 100 Cataulacus ebrardi Cataulacus hispidulus 98 Ocymyrmex fortior cf 100 Cardiocondyla mauritanica 100 Cardiocondyla MY01 Cardiocondyla thoracica 100 Monomorium robustior 100 Monomorium criniceps Monomorium destructor 100 100 Huberia brounii Podomyrma silvicola 100 Liomyrmex gestroi 100 100 Metapone madagascarica Metapone PG01 100 100 Eutetramorium mocquerysi Royidris notorthotenes 82 100 Royidris longiseta 100 Malagidris jugum 100 Malagidris belti Myrmisaraka brevis Vitsika suspicax Figure S1b 243 taxa

100 Apterogyna ZA01 Amblyopone australis 84 Stigmatomma pallipes 83 63 Proceratium stictum 70 Paraponera clavata 100 Platythyrea mocquerysi 100 Ankylomyrma coronacantha Tatuidris tatusia 100 Tatuidris tatusia_2 100 Aneuretus simoni 100 Dolichoderus pustulatus 100 Tetraponera rufonigra Myrmecia pyriformis 100 Nothomyrmecia macrops 100 Lasius californicus 100 Myrmelachista flavocotea Acanthoponera minor 100 Rhytidoponera chalybaea 100 100 Manica bradleyi 100 Manica rubida 100 Myrmica kotokui 100 Myrmica rugiventris 92 Myrmica striolagaster 100 Myrmica tahoensis 99 Pogonomyrmex angustus 100 Hylomyrma blandiens 100 Pogonomyrmex striatinodus 100 Pogonomyrmex imberbiculus Pogonomyrmex subdentatus 100 Pogonomyrmex vermiculatus 100 100 Aphaenogaster albisetosa 100 Messor andrei 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 100 Dolopomyrmex pilatus 100 Rogeria nevadensis Bariamyrma hispidula 100 100 Rogeria stigmatica 82 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 Solenopsis xyloni 100 Solenopsis molesta Solenopsis succinea 100 Megalomyrmex silvestrii 75 Tyrannomyrmex legatus 66 Monomorium brocha 57 Monomorium latinode 100 Monomorium sechellense 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 90 Tranopelta gilva 89 Diaphoromyrma sofiae 100 Lachnomyrmex scrobiculatus 100 Blepharidatta conops Allomerus octoarticulatus 100 Wasmannia auropunctata 95 71 Acanthognathus ocellatus 92 Daceton armigerum 100 Lenomyrmex colwelli 100 Microdaceton tibialis 100 Orectognathus versicolor 100 100 Colobostruma unicorna 92 Epopostruma monstrosa 100 Mesostruma browni 100 Apterostigma auriculatum 100 Mycocepurus goeldii 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 100 Trachymyrmex arizonensis 100 Procryptocerus scabriusculus 99 Cephalotes atratus Cephalotes texanus 100 Pheidole fimbriata 100 Pheidole rhea 100 Pheidole clydei 67 100 Pheidole hyatti 100 Pheidole lucioccipitalis 100 Pheidole pallidula 95 52 Anisopheidole antipodum Machomyrma dispar 100 Pheidole longispinosa 100 Protalaridris armata 91 Rhopalothrix isthmica 100 Basiceros manni 100 Octostruma EC01 81 100 100 Eurhopalothrix australis Eurhopalothrix gravis 100 Talaridris mandibularis Phalacromyrmex fugax 100 Pilotrochus besmerus 100 Strumigenys ambatrix 100 100 Strumigenys nitens Strumigenys gundlachi 100 100 Strumigenys biolleyi Strumigenys erynnes 100 100 Strumigenys ludovici 100 Strumigenys rogeri 87 Strumigenys coveri Strumigenys dicomas 100 Strumigenys maxillaris 95 Strumigenys olsoni 96 98 Strumigenys exiguaevitae 100 Strumigenys membranifera 92 Strumigenys ocypete 100 Strumigenys chiricahua 100 Strumigenys emmae Strumigenys chyzeri Strumigenys godeffroyi 100 Tetheamyrma subspongia 100 Paratopula TH01 59 Poecilomyrma myrmecodiae 100 Romblonella scrobifera 60 Rotastruma recava Rotastruma stenoceps 100 Gauromyrmex acanthinus 78 100 Vombisidris bilongrudi 99 Harpagoxenus sublaevis 82 Formicoxenus diversipilosus 56 100 Leptothorax muscorum complex 73 Temnothorax poeyi 100 Temnothorax rugatulus Chalepoxenus kutteri 100 100 Myrmoxenus bernardi 100 Rhopalomastix rothneyi 100 Terataner bottegoi 100 Terataner MG02 Terataner MG27 100 Kartidris sparsipila 99 Cyphoidris exalta 100 Dicroaspis KM01 100 100 Calyptomyrmex kaurus Myrmicine genus26 PH02 100 Vollenhovia emeryi 100 Tetramorium venator 100 Tetramorium validiusculum Tetramorium CF01 100 100 Tetramorium taylori 100 Tetramorium caespitum cf 86 100 Strongylognathus testaceus Anergates atratulus 100 Teleutomyrmex kutteri 97 Rhoptromyrmex transversinodis 100 Tetramorium MG125 Tetramorium severini 100 Tetramorium spinosum Dacatria templaris 100 Proatta butteli 100 Myrmecina PG01 Myrmecina americana 78 Myrmecina graminicola 87 Acanthomyrmex ferox 100 Mayriella ebbei 52 Propodilobus pingorum 96 Ancyridris polyrhachioides nr Lordomyrma bhutanensis cf 74 Lordomyrma desupra 93 Lophomyrmex ambiguus 65 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 Carebara affinis 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 Malagidris jugum 100 Malagidris belti 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.

Figure S2 18S

Apterogyna ZA01 Platythyrea mocquerysi Paraponera clavata 78 Amblyopone australis Stigmatomma pallipes 99 Proceratium stictum 99 Ankylomyrma coronacantha Tatuidris tatusia Tatuidris tatusia_2 97 Myrmecia pyriformis 98 80 Nothomyrmecia macrops 96 Tetraponera rufonigra 100 Aneuretus simoni Dolichoderus pustulatus Acanthoponera minor 80 Rhytidoponera chalybaea 81 Lasius californicus Myrmelachista flavocotea Hylomyrma blandiens Manica bradleyi Manica rubida Pogonomyrmex angustus Pogonomyrmex striatinodus 91 Pogonomyrmex imberbiculus Pogonomyrmex subdentatus Pogonomyrmex vermiculatus Myrmica kotokui 100 58 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 Atopomyrmex mocquerysi 58 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 Tetramorium venator 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 Kempfidris inusualis Mycetarotes acutus Mycetosoritis hartmanni Mycocepurus goeldii Myrmicaria carinata Myrmicaria exigua Malagidris jugum Malagidris belti 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 Wasmannia auropunctata 100 Acanthognathus ocellatus Microdaceton tibialis 99 Anillomyrma AFRC TZ 01 Anillomyrma decamera 92 Bariamyrma hispidula Mayriella ebbei 60 Daceton armigerum Kalathomyrmex emeryi 78 Dilobocondyla borneensis cf Secostruma MY01 51 Meranoplus castaneus Meranoplus radamae 100 Metapone madagascarica Metapone PG01 84 Monomorium hanneli 96 Rogeria nevadensis Royidris notorthotenes Royidris longiseta 68 Myrmicine genus26 PH02 Xenomyrmex floridanus 97 Ochetomyrmex neopolitus 99 Ochetomyrmex semipolitus Tranopelta subterranea Tranopelta gilva 99 Ancyridris polyrhachioides nr Lordomyrma bhutanensis cf Lordomyrma desupra 77 Cardiocondyla mauritanica Cardiocondyla MY01 Cardiocondyla thoracica 98 Eurhopalothrix australis 86 Eurhopalothrix gravis Talaridris mandibularis 58 Oxyopomyrmex santschii 100 Messor denticornis Messor wasmanni 100 Monomorium criniceps Monomorium destructor Monomorium robustior 100 Terataner bottegoi Terataner MG02 Terataner MG27 85 Monomorium latinode Monomorium sechellense 86 Adelomyrmex SC01 Oxyepoecus vezenyii Colobostruma unicorna 74 Epopostruma monstrosa Mesostruma browni Orectognathus versicolor 71 Diplomorium longipenne 55 Carebara vidua Carebara nosindambo 88 97 Carebara alperti Carebara AU01 97 Carebara urichi Carebara affinis Crematogaster acuta Crematogaster adrepens 90 61 Crematogaster modiglianii Crematogaster sordidula 68 Myrmecina americana Myrmecina graminicola Myrmecina PG01 Baracidris sitra 96 Solenopsis fugax Solenopsis xyloni Carebarella PA01 Solenopsis mameti 87 79 Solenopsis molesta Solenopsis papuana Solenopsis succinea Solenopsis ZA02 Monomorium exiguum 96 Monomorium madecassum Monomorium MG04 Monomorium pharaonis 89 Monomorium ergatogyna Monomorium fieldi nr Strumigenys ambatrix Strumigenys erynnes Strumigenys ludovici 98 Strumigenys biolleyi Strumigenys gundlachi 98 99 Strumigenys nitens Strumigenys membranifera Strumigenys godeffroyi Monomorium ferodens Strumigenys chiricahua Strumigenys exiguaevitae Strumigenys maxillaris 69 Strumigenys ocypete Strumigenys olsoni Strumigenys chyzeri Strumigenys emmae Strumigenys rogeri 100 Strumigenys coveri Strumigenys dicomas Figure S2 28S

Apterogyna ZA01 Paraponera clavata Proceratium stictum 98 Amblyopone australis Stigmatomma pallipes 100 100 Ankylomyrma coronacantha Tatuidris tatusia Tatuidris tatusia_2 Platythyrea mocquerysi 100 Acanthoponera minor Rhytidoponera chalybaea 51 100 Lasius californicus Myrmelachista flavocotea 100 Myrmecia pyriformis 84 Nothomyrmecia macrops 57 Tetraponera rufonigra 96 Aneuretus simoni Dolichoderus pustulatus Acanthognathus ocellatus Adelomyrmex paratristani 88 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 Xenomyrmex floridanus 100 Allomerus octoarticulatus Wasmannia auropunctata 100 Anillomyrma AFRC TZ 01 Anillomyrma decamera 54 Aphaenogaster albisetosa Lophomyrmex ambiguus 100 Atopomyrmex mocquerysi Myrmisaraka brevis 52 Austromorium flavigaster Monomorium brocha 59 Blepharidatta conops Chalepoxenus kutteri 100 Cataulacus ebrardi Cataulacus hispidulus 100 Crematogaster acuta Crematogaster sordidula 100 Dilobocondyla borneensis cf Secostruma MY01 91 Megalomyrmex silvestrii Monomorium denticulatum 99 Meranoplus castaneus 100 Meranoplus radamae Royidris notorthotenes Royidris longiseta 100 Myrmicaria carinata 99 Myrmicaria exigua Malagidris jugum Malagidris belti 100 Nesomyrmex echinatinodis Nesomyrmex wilda 100 Ochetomyrmex neopolitus Ochetomyrmex semipolitus 81 Ocymyrmex fortior cf 100 Phalacromyrmex fugax Tranopelta subterranea Tranopelta gilva 57 Acanthomyrmex ferox 100 Pristomyrmex orbiceps Pristomyrmex punctatus 98 Cardiocondyla MY01 58 Cardiocondyla mauritanica Cardiocondyla thoracica 98 Crematogaster modiglianii 90 Crematogaster adrepens Crematogaster emeryana 93 Eurhopalothrix australis Eurhopalothrix gravis 51 Talaridris mandibularis 73 Formicoxenus diversipilosus Harpagoxenus sublaevis Leptothorax muscorum complex 100 Liomyrmex gestroi 100 Metapone madagascarica Metapone PG01 100 Myrmecina PG01 99 Myrmecina americana Myrmecina graminicola 100 Nesomyrmex madecassus 72 Nesomyrmex MG01 Nesomyrmex MG08 100 Terataner bottegoi Terataner MG02 Terataner MG27 71 Paratopula TH01 Romblonella scrobifera 99 Rotastruma recava Rotastruma stenoceps 66 Dolopomyrmex pilatus 96 Monomorium latinode 100 Monomorium robustior 74 Monomorium criniceps Monomorium destructor 100 Pogonomyrmex angustus 90 Hylomyrma blandiens 100 Pogonomyrmex striatinodus 100 Pogonomyrmex imberbiculus Pogonomyrmex subdentatus Pogonomyrmex vermiculatus 100 Manica bradleyi 100 Manica rubida 100 Myrmica kotokui 53 Myrmica rugiventris 78 Myrmica striolagaster Myrmica tahoensis Atta texana 100 Cyphomyrmex cornutus Mycetarotes acutus 63 Cyphomyrmex longiscapus cf 60 Mycetosoritis hartmanni 72 Acromyrmex versicolor Trachymyrmex arizonensis Daceton armigerum 98 Lenomyrmex colwelli 63 Microdaceton tibialis Colobostruma unicorna 100 Epopostruma monstrosa Mesostruma browni Orectognathus versicolor Oxyopomyrmex santschii 54 98 Aphaenogaster swammerdami 100 Aphaenogaster pythia Stenamma expolitum 52 Aphaenogaster araneoides 70 Aphaenogaster occidentalis 84 Messor denticornis Messor wasmanni 80 Carebara nosindambo 73 Stegomyrmex manni 100 Diplomorium longipenne 100 Carebara vidua 100 Carebara urichi 75 Carebara affinis 100 Carebara alperti Carebara AU01 58 84 Solenopsis succinea 83 Solenopsis mameti Solenopsis molesta 88 Solenopsis ZA02 85 Solenopsis fugax 100 Solenopsis xyloni Kempfidris inusualis Tropidomyrmex elianae Anisopheidole antipodum Machomyrma dispar Pheidole clydei 99 Pheidole fimbriata Pheidole longispinosa Pheidole pallidula Pheidole rhea 97 Pheidole hyatti Pheidole lucioccipitalis Bariamyrma hispidula 57 Monomorium fisheri Monomorium hanneli 96 Monomorium sechellense 100 Monomorium exiguum Monomorium fieldi nr 92 Monomorium MG04 Monomorium pharaonis 99 Monomorium ergatogyna Monomorium madecassum Tetramorium venator Rhoptromyrmex transversinodis Tetramorium caespitum cf 100 97 Tetramorium taylori 61 Tetramorium severini Tetramorium spinosum Strongylognathus testaceus 54 Tetramorium CF01 Tetramorium MG125 Tetramorium validiusculum 99 Anergates atratulus Teleutomyrmex kutteri Strumigenys ambatrix Strumigenys exiguaevitae Strumigenys olsoni Strumigenys emmae 100 Strumigenys coveri 100 Strumigenys dicomas 87 Strumigenys rogeri Strumigenys chiricahua Strumigenys ocypete 65 Strumigenys membranifera 51 72 63 Strumigenys godeffroyi Strumigenys nitens Strumigenys chyzeri 98 Strumigenys maxillaris 54 Strumigenys erynnes 100 Strumigenys ludovici Strumigenys gundlachi Strumigenys biolleyi Figure S2 Abd-A

Apterogyna ZA01 Platythyrea mocquerysi Paraponera clavata 77 Ankylomyrma coronacantha 81 100 Proceratium stictum 70 Tatuidris tatusia Tatuidris tatusia_2 Stigmatomma pallipes Amblyopone australis 81 Tetraponera rufonigra 96 Acanthoponera minor Rhytidoponera chalybaea 85 Aneuretus simoni 100 Dolichoderus pustulatus 74 Lasius californicus 77 Myrmelachista flavocotea 98 Myrmecia pyriformis Nothomyrmecia macrops 100 Hylomyrma blandiens 65 Pogonomyrmex angustus Pogonomyrmex striatinodus 100 Pogonomyrmex imberbiculus 87 Pogonomyrmex subdentatus 89 Pogonomyrmex vermiculatus 100 Manica bradleyi 100 Manica rubida 90 Myrmica kotokui 91 Myrmica rugiventris 100 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 Carebara affinis Podomyrma silvicola Pristomyrmex orbiceps Proatta butteli Stegomyrmex manni Tetheamyrma subspongia Xenomyrmex floridanus 66 Adelomyrmex paratristani Baracidris sitra 57 Adlerzia froggatti Recurvidris TH01 100 Anillomyrma AFRC TZ 01 Anillomyrma decamera 89 Carebarella PA01 Solenopsis papuana 100 Dilobocondyla borneensis cf Secostruma MY01 100 Melissotarsus insularis Rhopalomastix rothneyi 91 100 Meranoplus castaneus Meranoplus radamae 66 Monomorium brocha Monomorium sechellense 95 Monomorium denticulatum Oxyepoecus vezenyii 96 Monomorium hanneli 100 Tyrannomyrmex legatus Kempfidris inusualis 100 Tropidomyrmex elianae Royidris notorthotenes Royidris longiseta 100 Myrmicaria carinata 100 Myrmicaria exigua Malagidris jugum Malagidris belti 61 Rogeria nevadensis Rogeria stigmatica 75 Monomorium fisheri 91 Adelomyrmex SC01 Cryptomyrmex boltoni 100 Cardiocondyla MY01 87 Cardiocondyla mauritanica Cardiocondyla thoracica 82 Ocymyrmex fortior cf 100 Cataulacus ebrardi Cataulacus hispidulus 70 Eutetramorium mocquerysi Huberia brounii Perissomyrmex snyderi 83 Liomyrmex gestroi 99 Metapone madagascarica Metapone PG01 100 Monomorium robustior 100 Monomorium criniceps Monomorium destructor 100 Myrmecina PG01 65 Myrmecina americana Myrmecina graminicola 99 Ancyridris polyrhachioides nr Propodilobus pingorum 100 Lordomyrma bhutanensis cf Lordomyrma desupra 55 Carebara nosindambo Pristomyrmex punctatus 100 Carebara alperti Carebara AU01 100 Nesomyrmex echinatinodis 77 Nesomyrmex wilda 100 Nesomyrmex madecassus Nesomyrmex MG01 91 Nesomyrmex MG08 90 Acanthomyrmex ferox 100 Crematogaster acuta 96 Crematogaster sordidula 99 Crematogaster modiglianii 100 Crematogaster adrepens Crematogaster emeryana 100 Monomorium fieldi nr 93 Monomorium ergatogyna 58 Monomorium madecassum 95 Monomorium pharaonis 100 Monomorium exiguum Monomorium MG04 Solenopsis mameti 100 Solenopsis xyloni 100 Solenopsis fugax Solenopsis ZA02 77 Solenopsis molesta Solenopsis succinea 71 Paratopula TH01 100 Rotastruma recava 73 Rotastruma stenoceps 61 Poecilomyrma myrmecodiae 73 Romblonella scrobifera 100 Terataner bottegoi 92 Terataner MG02 Terataner MG27 100 Gauromyrmex acanthinus Vombisidris bilongrudi 59 100 Harpagoxenus sublaevis 71 Formicoxenus diversipilosus 79 Leptothorax muscorum complex 97 Myrmoxenus bernardi Temnothorax rugatulus 93 Chalepoxenus kutteri Temnothorax poeyi Stenamma striatulum 100 Aphaenogaster albisetosa Messor andrei 100 75 Stenamma dyscheres Stenamma expolitum 100 Aphaenogaster pythia Aphaenogaster swammerdami 97 100 Goniomma blanci Oxyopomyrmex santschii 99 Aphaenogaster araneoides 100 Aphaenogaster occidentalis 87 Messor denticornis Messor wasmanni 98 Calyptomyrmex kaurus 97 Myrmicine genus26 PH02 Vollenhovia emeryi 86 Rhoptromyrmex transversinodis Tetramorium MG125 96 Tetramorium severini 100 Tetramorium spinosum Anergates atratulus 100 Strongylognathus testaceus Teleutomyrmex kutteri Tetramorium caespitum cf 63 Tetramorium taylori 99 100 Tetramorium CF01 Tetramorium venator Tetramorium validiusculum Blepharidatta conops Diaphoromyrma sofiae Lachnomyrmex scrobiculatus Octostruma EC01 Procryptocerus scabriusculus 100 Allomerus octoarticulatus Wasmannia auropunctata 100 Cephalotes atratus Cephalotes texanus 94 Phalacromyrmex fugax Pilotrochus besmerus 64 Basiceros manni 86 Protalaridris armata Rhopalothrix isthmica 69 Eurhopalothrix australis 100 Eurhopalothrix gravis 100 Talaridris mandibularis 94 Ochetomyrmex neopolitus 100 Ochetomyrmex semipolitus Tranopelta subterranea Tranopelta gilva Pheidole fimbriata 69 Pheidole rhea Pheidole hyatti Pheidole longispinosa 99 75 Pheidole pallidula 63 Anisopheidole antipodum Pheidole clydei 76 Machomyrma dispar 99 Pheidole lucioccipitalis Strumigenys ambatrix 100 Strumigenys gundlachi Strumigenys biolleyi 83 Strumigenys nitens Strumigenys chiricahua 73 Strumigenys erynnes Strumigenys exiguaevitae Strumigenys ludovici Strumigenys maxillaris 92 Strumigenys membranifera Strumigenys ocypete Strumigenys olsoni 86 Strumigenys chyzeri Strumigenys emmae Strumigenys godeffroyi 100 Strumigenys rogeri 100 Strumigenys coveri Strumigenys dicomas Acanthognathus ocellatus Daceton armigerum Kalathomyrmex emeryi Lenomyrmex colwelli Microdaceton tibialis Myrmicocrypta infuscata cf 58 Orectognathus versicolor 94 Apterostigma auriculatum Mycocepurus goeldii 73 Colobostruma unicorna 98 Epopostruma monstrosa Mesostruma browni 92 Mycetarotes acutus 69 Mycetosoritis hartmanni 78 Cyphomyrmex cornutus 76 Cyphomyrmex longiscapus cf 100 Acromyrmex versicolor Atta texana Trachymyrmex arizonensis Figure S2 EF1aF1

Apterogyna ZA01 Myrmecia pyriformis Amblyopone australis 100 Platythyrea mocquerysi Proceratium stictum 77 Stigmatomma pallipes 100 100 Ankylomyrma coronacantha Tatuidris tatusia Tatuidris tatusia_2 Nothomyrmecia macrops 72 Paraponera clavata 98 Rhytidoponera chalybaea Tetraponera rufonigra 75 Acanthoponera minor 74 Dolichoderus pustulatus 100 Lasius californicus Myrmelachista flavocotea Aneuretus simoni Anillomyrma AFRC TZ 01 Anillomyrma decamera 65 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 Solenopsis papuana 55 Adelomyrmex paratristani Tyrannomyrmex legatus 56 Megalomyrmex silvestrii 96 Rogeria stigmatica Kempfidris inusualis Tropidomyrmex elianae 100 Myrmicaria carinata Myrmicaria exigua 100 Cardiocondyla mauritanica 94 Cardiocondyla MY01 Cardiocondyla thoracica 61 Monomorium ferodens Monomorium fisheri 99 Cryptomyrmex boltoni 72 Adelomyrmex SC01 Baracidris sitra 77 Hylomyrma blandiens 82 Pogonomyrmex striatinodus 79 100 Pogonomyrmex imberbiculus 54 Pogonomyrmex subdentatus Pogonomyrmex vermiculatus Monomorium exiguum 98 Monomorium fieldi nr Monomorium MG04 56 Monomorium ergatogyna Monomorium madecassum Monomorium pharaonis 100 Solenopsis fugax 100 Solenopsis ZA02 100 Solenopsis mameti 99 Solenopsis molesta Solenopsis succinea Solenopsis xyloni Acanthognathus ocellatus 57 56 Manica bradleyi 68 Manica rubida Myrmica kotokui 100 Myrmica rugiventris Myrmica striolagaster Myrmica tahoensis Calyptomyrmex kaurus 100 Cataulacus ebrardi 51 Cataulacus hispidulus 95 53 Nesomyrmex MG01 56 Nesomyrmex madecassus Nesomyrmex MG08 90 Terataner bottegoi Terataner MG02 Terataner MG27 Adlerzia froggatti Aphaenogaster araneoides Carebara vidua Cyphoidris exalta Dacatria templaris 67 Dacetinops ignotus cf Diaphoromyrma sofiae Dicroaspis KM01 Eutetramorium mocquerysi Gauromyrmex acanthinus Huberia brounii Kartidris sparsipila Mayriella ebbei Melissotarsus insularis Malagidris jugum Malagidris belti Myrmisaraka brevis Vitsika suspicax Myrmicine genus26 PH02 Pheidole fimbriata Proatta butteli Procryptocerus scabriusculus Recurvidris TH01 Rhopalomastix rothneyi Stegomyrmex manni Tetheamyrma subspongia Vollenhovia emeryi Vombisidris bilongrudi 83 Acanthomyrmex ferox Diplomorium longipenne 100 Aphaenogaster albisetosa Messor andrei 65 Aphaenogaster pythia Aphaenogaster swammerdami 89 Carebara urichi Carebara affinis 100 Dilobocondyla borneensis cf Secostruma MY01 100 Meranoplus castaneus 100 Meranoplus radamae Royidris notorthotenes Royidris longiseta 100 Nesomyrmex echinatinodis Nesomyrmex wilda 77 Perissomyrmex snyderi Podomyrma silvicola 99 Pristomyrmex orbiceps Pristomyrmex punctatus 97 Carebara nosindambo 57 Carebara alperti Carebara AU01 100 Formicoxenus diversipilosus Harpagoxenus sublaevis Leptothorax muscorum complex 72 Liomyrmex gestroi 85 Metapone madagascarica Metapone PG01 97 Monomorium criniceps Monomorium destructor Monomorium robustior 98 Stenamma expolitum 98 Stenamma dyscheres Stenamma striatulum 66 Allomerus octoarticulatus 60 Wasmannia auropunctata 60 Blepharidatta conops Lachnomyrmex scrobiculatus Chalepoxenus kutteri 96 Myrmoxenus bernardi Temnothorax poeyi Temnothorax rugatulus 70 Poecilomyrma myrmecodiae Romblonella scrobifera 100 Rotastruma recava Rotastruma stenoceps 57 54 Lophomyrmex ambiguus 99 Ancyridris polyrhachioides nr Propodilobus pingorum 91 Lordomyrma bhutanensis cf 100 Lordomyrma desupra 99 Goniomma blanci Oxyopomyrmex santschii 87 Aphaenogaster occidentalis 100 Messor denticornis Messor wasmanni 58 Ocymyrmex fortior cf 51 Paratopula TH01 100 Myrmecina PG01 97 Myrmecina americana Myrmecina graminicola Atopomyrmex mocquerysi 82 100 Crematogaster acuta Crematogaster sordidula 77 Crematogaster modiglianii 99 Crematogaster adrepens Crematogaster emeryana 100 Cephalotes atratus 80 100 Cephalotes texanus 100 Ochetomyrmex neopolitus 100 Ochetomyrmex semipolitus Tranopelta subterranea Tranopelta gilva Basiceros manni 98 Octostruma EC01 Protalaridris armata Rhopalothrix isthmica 95 Eurhopalothrix australis 97 Eurhopalothrix gravis Talaridris mandibularis 82 Daceton armigerum Lenomyrmex colwelli 65 Microdaceton tibialis 100 Orectognathus versicolor 94 Colobostruma unicorna Epopostruma monstrosa Mesostruma browni Pheidole clydei 96 Pheidole hyatti 67 Pheidole rhea Anisopheidole antipodum Machomyrma dispar 59 Pheidole longispinosa Pheidole lucioccipitalis Pheidole pallidula Xenomyrmex floridanus 82 Apterostigma auriculatum 64 Myrmicocrypta infuscata cf 54 Mycocepurus goeldii 96 Kalathomyrmex emeryi 51 Mycetarotes acutus Mycetosoritis hartmanni 99 Cyphomyrmex longiscapus cf 51 Cyphomyrmex cornutus 100 Trachymyrmex arizonensis 61 Acromyrmex versicolor Atta texana Tetramorium venator Rhoptromyrmex transversinodis Tetramorium validiusculum 100 Tetramorium CF01 73 Tetramorium taylori 73 Tetramorium MG125 73 Tetramorium severini Tetramorium spinosum 97 Anergates atratulus 67 Teleutomyrmex kutteri 100 Strongylognathus testaceus Tetramorium caespitum cf 98 Phalacromyrmex fugax Pilotrochus besmerus Strumigenys ambatrix 99 95 Strumigenys rogeri Strumigenys erynnes Strumigenys ludovici 69 Strumigenys coveri 93 Strumigenys dicomas 82 Strumigenys gundlachi Strumigenys nitens Strumigenys biolleyi Strumigenys exiguaevitae Strumigenys olsoni 78 Strumigenys godeffroyi 82 75 Strumigenys chiricahua Strumigenys membranifera Strumigenys ocypete 61 Strumigenys maxillaris Strumigenys chyzeri Strumigenys emmae Figure S2 LW Rh

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 Vombisidris bilongrudi 96 Dacatria templaris Proatta butteli 99 Huberia brounii Podomyrma silvicola 100 Meranoplus castaneus Meranoplus radamae 83 Myrmicine genus26 PH02 Vollenhovia emeryi 100 Pristomyrmex orbiceps Pristomyrmex punctatus 100 Blepharidatta conops 100 Allomerus octoarticulatus Wasmannia auropunctata 87 Apterostigma auriculatum 99 Mycocepurus goeldii Myrmicocrypta infuscata cf 98 Colobostruma unicorna 100 Epopostruma monstrosa Mesostruma browni 68 Gauromyrmex acanthinus Mayriella ebbei Tetheamyrma subspongia 72 Lophomyrmex ambiguus 99 Melissotarsus insularis Rhopalomastix rothneyi 100 Myrmecina PG01 100 Myrmecina americana Myrmecina graminicola 100 Terataner MG02 54 Terataner bottegoi Terataner MG27 100 Lordomyrma bhutanensis cf 77 Lordomyrma desupra 51 Ancyridris polyrhachioides nr Propodilobus pingorum 100 Temnothorax poeyi 56 Temnothorax rugatulus 91 Chalepoxenus kutteri 100 Myrmoxenus bernardi 100 Ochetomyrmex neopolitus 98 Ochetomyrmex semipolitus Tranopelta subterranea Tranopelta gilva 100 Cephalotes atratus 90 Cephalotes texanus 97 Procryptocerus scabriusculus 100 Dilobocondyla borneensis cf Secostruma MY01 97 60 Eutetramorium mocquerysi 100 Malagidris jugum 100 Malagidris belti Myrmisaraka brevis Vitsika suspicax 78 Paratopula TH01 92 Poecilomyrma myrmecodiae 56 Romblonella scrobifera 100 Rotastruma recava Rotastruma stenoceps 99 Carebara vidua 100 Carebara nosindambo 100 Carebara alperti 53 Carebara AU01 100 Carebara urichi Carebara affinis Crematogaster modiglianii 57 Ocymyrmex fortior cf 96 Crematogaster acuta Crematogaster sordidula 71 Crematogaster adrepens Crematogaster emeryana 95 Kalathomyrmex emeryi 78 Mycetarotes acutus 90 Mycetosoritis hartmanni 100 Cyphomyrmex cornutus 96 Cyphomyrmex longiscapus cf 100 Trachymyrmex arizonensis 92 Acromyrmex versicolor Atta texana 100 Pheidole rhea Pheidole fimbriata 57 Pheidole longispinosa 100 Pheidole lucioccipitalis 53 Anisopheidole antipodum Machomyrma dispar 70 Pheidole pallidula 94 Pheidole clydei Pheidole hyatti Basiceros manni Octostruma EC01 58 Protalaridris armata 56 Rhopalothrix isthmica 100 Cataulacus ebrardi Cataulacus hispidulus 91 Eurhopalothrix australis 99 Eurhopalothrix gravis Talaridris mandibularis 99 Xenomyrmex floridanus 100 Formicoxenus diversipilosus 94 Harpagoxenus sublaevis Leptothorax muscorum complex 100 Nesomyrmex echinatinodis 80 Nesomyrmex wilda 100 Nesomyrmex MG08 88 Nesomyrmex madecassus Nesomyrmex MG01 68 Liomyrmex gestroi 100 Metapone madagascarica 70 Metapone PG01 100 Monomorium destructor 75 Monomorium criniceps 54 100 Monomorium robustior 61 Royidris notorthotenes Royidris longiseta 100 Cardiocondyla mauritanica 100 Cardiocondyla MY01 Cardiocondyla thoracica 93 Tetramorium spinosum Tetramorium venator Tetramorium validiusculum 100 60 Rhoptromyrmex transversinodis Tetramorium MG125 100 Tetramorium CF01 Tetramorium taylori 70 Tetramorium severini 100 Tetramorium caespitum cf 94 Strongylognathus testaceus 99 Anergates atratulus Teleutomyrmex kutteri 100 Aphaenogaster albisetosa 99 Messor andrei 100 Stenamma expolitum 59 Stenamma dyscheres 100 Stenamma striatulum 72 Aphaenogaster araneoides 100 Aphaenogaster pythia 97 100 Aphaenogaster swammerdami 100 Goniomma blanci Oxyopomyrmex santschii 100 Aphaenogaster occidentalis 100 Messor denticornis Messor wasmanni 99 Phalacromyrmex fugax Pilotrochus besmerus 61 Strumigenys ambatrix Strumigenys erynnes 97 Strumigenys ludovici 92 Strumigenys biolleyi Strumigenys gundlachi 73 Strumigenys nitens 80 Strumigenys exiguaevitae Strumigenys maxillaris 100 Strumigenys coveri Strumigenys dicomas Strumigenys rogeri 56 Strumigenys olsoni Strumigenys chiricahua 100 Strumigenys membranifera Strumigenys ocypete Strumigenys godeffroyi 66 Strumigenys chyzeri Strumigenys emmae 99 Hylomyrma blandiens Pogonomyrmex angustus 93 Pogonomyrmex striatinodus 100 Pogonomyrmex imberbiculus 93 Pogonomyrmex subdentatus Pogonomyrmex vermiculatus 56 100 Manica bradleyi 100 Manica rubida 100 Myrmica kotokui 98 Myrmica rugiventris 84 Myrmica striolagaster 63 Myrmica tahoensis Aneuretus simoni 100 Acanthoponera minor Rhytidoponera chalybaea 98 Lasius californicus 64 Myrmelachista flavocotea 71 Dolichoderus pustulatus 92 Tetraponera rufonigra 100 Myrmecia pyriformis Nothomyrmecia macrops 61 Platythyrea mocquerysi Stigmatomma pallipes 57 56 Amblyopone australis Proceratium stictum 70 Paraponera clavata 100 100 Ankylomyrma coronacantha Tatuidris tatusia Tatuidris tatusia_2 Austromorium flavigaster Bariamyrma hispidula Monomorium sechellense Oxyepoecus vezenyii 61 Megalomyrmex silvestrii Monomorium denticulatum 70 Monomorium antarcticum 57 Monomorium fisheri Rogeria nevadensis 87 Adelomyrmex paratristani 100 Baracidris sitra 98 Adelomyrmex SC01 90 Cryptomyrmex boltoni 100 Carebarella PA01 100 Solenopsis papuana 99 Solenopsis fugax 100 Solenopsis ZA02 93 Solenopsis xyloni 99 Solenopsis mameti 100 Solenopsis molesta Solenopsis succinea Monomorium ferodens Monomorium hanneli 86 Monomorium brocha 58 Monomorium latinode 51 59 Tyrannomyrmex legatus 100 Anillomyrma AFRC TZ 01 89 Anillomyrma decamera 100 Myrmicaria carinata 100 Myrmicaria exigua Kempfidris inusualis 53 Tropidomyrmex elianae 99 Dolopomyrmex pilatus Monomorium ergatogyna 100 Monomorium madecassum Monomorium MG04 63 Monomorium fieldi nr 58 Monomorium exiguum Monomorium pharaonis Figure S2 ArgK

Apterogyna ZA01 Platythyrea mocquerysi 100 100 Amblyopone australis Stigmatomma pallipes 93 Paraponera clavata 57 Proceratium stictum 100 100 Ankylomyrma coronacantha Tatuidris tatusia Tatuidris tatusia_2 92 Acanthoponera minor Rhytidoponera chalybaea 100 Lasius californicus Myrmelachista flavocotea 84 Aneuretus simoni Dolichoderus pustulatus 85 Tetraponera rufonigra 100 Myrmecia pyriformis Nothomyrmecia macrops 100 Manica bradleyi 100 Manica rubida 100 Myrmica kotokui Myrmica rugiventris 100 Myrmica striolagaster Myrmica tahoensis 100 Hylomyrma blandiens Pogonomyrmex angustus 86 66 Pogonomyrmex striatinodus 100 Pogonomyrmex imberbiculus 55 Pogonomyrmex subdentatus Pogonomyrmex vermiculatus Stenamma dyscheres 100 Aphaenogaster albisetosa 100 Messor andrei 70 Stenamma expolitum Stenamma striatulum 100 Aphaenogaster pythia 99 Aphaenogaster swammerdami 85 100 Aphaenogaster araneoides 98 Goniomma blanci Oxyopomyrmex santschii 100 Aphaenogaster occidentalis 100 Messor denticornis Messor wasmanni 98 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 Xenomyrmex floridanus 100 Melissotarsus insularis Rhopalomastix rothneyi 100 Meranoplus castaneus 100 Meranoplus radamae Royidris notorthotenes Royidris longiseta 78 Myrmicine genus26 PH02 Vollenhovia emeryi 76 100 Liomyrmex gestroi 90 Metapone madagascarica Metapone PG01 100 Monomorium robustior 71 Monomorium criniceps Monomorium destructor 100 Myrmecina PG01 81 Myrmecina americana Myrmecina graminicola 68 Perissomyrmex snyderi 100 Pristomyrmex orbiceps Pristomyrmex punctatus 100 Terataner bottegoi 99 Terataner MG02 Terataner MG27 99 Lordomyrma desupra Propodilobus pingorum 79 Ancyridris polyrhachioides nr 100 Lordomyrma bhutanensis cf 89 Malagidris jugum 94 Malagidris belti 96 Myrmisaraka brevis Vitsika suspicax 100 Crematogaster acuta 100 Crematogaster sordidula 100 Crematogaster modiglianii 100 Crematogaster adrepens Crematogaster emeryana 100 Nesomyrmex echinatinodis 100 Nesomyrmex wilda 100 Nesomyrmex madecassus 79 Nesomyrmex MG01 Nesomyrmex MG08 71 Recurvidris TH01 92 Diplomorium longipenne 99 100 Carebara vidua 84 Carebara urichi Carebara affinis 65 Carebara nosindambo 100 Carebara alperti Carebara AU01 100 Cataulacus ebrardi Cataulacus hispidulus 98 94 98 Lophomyrmex ambiguus 100 Dilobocondyla borneensis cf Secostruma MY01 98 Paratopula TH01 100 Poecilomyrma myrmecodiae Romblonella scrobifera Rotastruma stenoceps 73 Atopomyrmex mocquerysi 100 Harpagoxenus sublaevis 72 93 Formicoxenus diversipilosus Leptothorax muscorum complex 88 Gauromyrmex acanthinus 88 Vombisidris bilongrudi 100 Chalepoxenus kutteri 95 Myrmoxenus bernardi 90 Temnothorax poeyi Temnothorax rugatulus Calyptomyrmex kaurus Cyphoidris exalta Dicroaspis KM01 100 93 Cardiocondyla mauritanica 58 Cardiocondyla MY01 Cardiocondyla thoracica 100 Tetramorium CF01 91 Tetramorium taylori 100 Tetramorium caespitum cf 89 Strongylognathus testaceus 100 93 Anergates atratulus 86 Teleutomyrmex kutteri Tetramorium venator 60 Tetramorium validiusculum 57 Tetramorium severini Tetramorium spinosum 89 Rhoptromyrmex transversinodis Tetramorium MG125 Bariamyrma hispidula Dolopomyrmex pilatus Megalomyrmex silvestrii Monomorium antarcticum Monomorium denticulatum Monomorium sechellense Oxyepoecus vezenyii Rogeria nevadensis Rogeria stigmatica Stegomyrmex manni Tyrannomyrmex legatus 100 Anillomyrma AFRC TZ 01 Anillomyrma decamera 93 Austromorium flavigaster Monomorium brocha 69 Monomorium ferodens Monomorium fisheri 99 51 Monomorium hanneli 100 Monomorium latinode Kempfidris inusualis Tropidomyrmex elianae 100 Myrmicaria carinata Myrmicaria exigua 73 Adelomyrmex paratristani 53 Baracidris sitra 51 Adelomyrmex SC01 Cryptomyrmex boltoni Monomorium ergatogyna 100 Monomorium pharaonis 58 Monomorium fieldi nr Monomorium madecassum 100 Monomorium exiguum Monomorium MG04 100 Carebarella PA01 100 Solenopsis papuana 100 Solenopsis fugax 100 Solenopsis ZA02 87 Solenopsis mameti 100 Solenopsis xyloni 83 Solenopsis molesta Solenopsis succinea 57 Acanthognathus ocellatus Blepharidatta conops Diaphoromyrma sofiae Lachnomyrmex scrobiculatus Lenomyrmex colwelli 100 Allomerus octoarticulatus Wasmannia auropunctata 100 Apterostigma auriculatum 76 Mycocepurus goeldii Myrmicocrypta infuscata cf 100 Procryptocerus scabriusculus 100 Cephalotes atratus Cephalotes texanus 100 Orectognathus versicolor 100 Colobostruma unicorna 100 Epopostruma monstrosa 100 Mesostruma browni 79 Ochetomyrmex neopolitus 100 Ochetomyrmex semipolitus Tranopelta subterranea Tranopelta gilva 100 Eurhopalothrix australis 100 Eurhopalothrix gravis 88 Talaridris mandibularis 99 89 Basiceros manni 96 Octostruma EC01 95 Protalaridris armata Rhopalothrix isthmica 65 Daceton armigerum 68 Kalathomyrmex emeryi Mycetosoritis hartmanni 100 77 Cyphomyrmex cornutus Mycetarotes acutus 66 Cyphomyrmex longiscapus cf 100 Acromyrmex versicolor Atta texana Trachymyrmex arizonensis 61 Microdaceton tibialis 99 Pheidole rhea 61 Pheidole fimbriata Anisopheidole antipodum 100 Pheidole longispinosa 87 Machomyrma dispar Pheidole lucioccipitalis 97 Pheidole pallidula 56 Pheidole clydei Pheidole hyatti 100 Phalacromyrmex fugax Pilotrochus besmerus 56 100 58 Strumigenys gundlachi Strumigenys nitens Strumigenys biolleyi 100 100 Strumigenys ambatrix Strumigenys erynnes Strumigenys ludovici 55 79 Strumigenys rogeri 88 Strumigenys coveri 93 Strumigenys dicomas Strumigenys maxillaris 100 Strumigenys olsoni 99 Strumigenys exiguaevitae 91 Strumigenys membranifera 75 Strumigenys chiricahua Strumigenys ocypete 87 Strumigenys emmae 100 Strumigenys chyzeri Strumigenys godeffroyi Figure S2 Top1

Apterogyna ZA01 85 Acanthoponera minor 100 Myrmecia pyriformis Nothomyrmecia macrops 78 Tetraponera rufonigra 94 Aneuretus simoni Dolichoderus pustulatus Proceratium stictum 100 83 Amblyopone australis 76 Stigmatomma pallipes 95 Platythyrea mocquerysi 66 Paraponera clavata 100 100 Ankylomyrma coronacantha Tatuidris tatusia 93 Tatuidris tatusia_2 100 Lasius californicus Myrmelachista flavocotea 100 Manica bradleyi 100 Manica rubida 100 Myrmica kotokui 84 Myrmica rugiventris 100 Myrmica striolagaster Myrmica tahoensis 100 Pogonomyrmex angustus 97 Hylomyrma blandiens 100 Pogonomyrmex striatinodus 86 97 Pogonomyrmex imberbiculus 54 Pogonomyrmex subdentatus Pogonomyrmex vermiculatus Rhytidoponera chalybaea Aphaenogaster araneoides 100 Aphaenogaster albisetosa 100 99 Messor andrei 63 100 Stenamma expolitum 92 Stenamma dyscheres 54 Stenamma striatulum 100 Aphaenogaster pythia 68 100 Aphaenogaster swammerdami 100 Goniomma blanci Oxyopomyrmex santschii 100 Aphaenogaster occidentalis 100 Messor denticornis Messor wasmanni Microdaceton tibialis Myrmicine genus26 PH02 100 Protalaridris armata 61 Rhopalothrix isthmica 67 74 Blepharidatta conops 90 Tetheamyrma subspongia 100 Allomerus octoarticulatus Wasmannia auropunctata Vollenhovia emeryi 51 Calyptomyrmex kaurus 100 Tetramorium CF01 93 Tetramorium taylori 100 Rhoptromyrmex transversinodis 100 Tetramorium MG125 100 Tetramorium severini 100 Tetramorium spinosum Tetramorium venator 88 Tetramorium validiusculum 100 Tetramorium caespitum cf 80 Strongylognathus testaceus 99 Anergates atratulus 96 Teleutomyrmex kutteri 93 Dolopomyrmex pilatus Rogeria stigmatica 92 Bariamyrma hispidula 100 Rogeria nevadensis 100 85 Austromorium flavigaster 99 Monomorium antarcticum Monomorium denticulatum 50 100 Oxyepoecus vezenyii 67 Kempfidris inusualis 71 Tropidomyrmex elianae 100 Carebarella PA01 100 Solenopsis papuana 100 Solenopsis fugax 100 Solenopsis ZA02 100 100 Solenopsis mameti 100 Solenopsis xyloni 81 Solenopsis molesta Solenopsis succinea Megalomyrmex silvestrii 92 Baracidris sitra 99 Adelomyrmex SC01 71 85 Cryptomyrmex boltoni 99 Adelomyrmex paratristani 100 Monomorium madecassum 100 Monomorium exiguum 97 Monomorium MG04 100 51 Monomorium ergatogyna 56 Monomorium fieldi nr Monomorium pharaonis 66 Monomorium ferodens Monomorium fisheri 86 Monomorium hanneli 94 Monomorium sechellense 100 Myrmicaria carinata 93 74 Myrmicaria exigua Monomorium brocha 96 Monomorium latinode Tyrannomyrmex legatus 100 Anillomyrma AFRC TZ 01 100 Anillomyrma decamera 100 Ochetomyrmex neopolitus 100 Ochetomyrmex semipolitus Tranopelta subterranea Tranopelta gilva 61 100 Octostruma EC01 97 Basiceros manni 99 Eurhopalothrix australis 100 100 Eurhopalothrix gravis Talaridris mandibularis 100 Phalacromyrmex fugax 100 Pilotrochus besmerus Strumigenys ambatrix 100 98 100 Strumigenys nitens Strumigenys gundlachi 98 Strumigenys biolleyi 100 Strumigenys rogeri 100 Strumigenys coveri 100 100 Strumigenys dicomas Strumigenys erynnes 82 Strumigenys ludovici Strumigenys maxillaris 100 Strumigenys chiricahua Strumigenys membranifera 52 61 Strumigenys ocypete Strumigenys exiguaevitae Strumigenys olsoni 78 Strumigenys godeffroyi 93 Strumigenys chyzeri Strumigenys emmae Lachnomyrmex scrobiculatus Stegomyrmex manni 97 Procryptocerus scabriusculus 100 Cephalotes atratus Cephalotes texanus 55 100 Pheidole fimbriata 100 Pheidole rhea Pheidole hyatti 100 Anisopheidole antipodum 76 Machomyrma dispar Pheidole clydei 77 Pheidole longispinosa 88 Pheidole lucioccipitalis Pheidole pallidula Acanthognathus ocellatus Daceton armigerum 98 Diaphoromyrma sofiae Lenomyrmex colwelli 100 Orectognathus versicolor 52 100 Colobostruma unicorna 100 Epopostruma monstrosa Mesostruma browni 63 100 Mycocepurus goeldii 100 Myrmicocrypta infuscata cf 96 Apterostigma auriculatum 99 Kalathomyrmex emeryi 100 Mycetarotes acutus 72 Mycetosoritis hartmanni 96 Cyphomyrmex cornutus 100 Cyphomyrmex longiscapus cf 100 Atta texana 61 Acromyrmex versicolor Trachymyrmex arizonensis Adlerzia froggatti Dacetinops ignotus cf Mayriella ebbei Perissomyrmex snyderi 52 Cyphoidris exalta Dicroaspis KM01 95 Dacatria templaris Proatta butteli 100 Dilobocondyla borneensis cf Secostruma MY01 58 Kartidris sparsipila Lophomyrmex ambiguus 100 Melissotarsus insularis Rhopalomastix rothneyi 100 Monomorium robustior 100 Monomorium criniceps Monomorium destructor 100 Ancyridris polyrhachioides nr 77 Lordomyrma desupra 55 Lordomyrma bhutanensis cf Propodilobus pingorum 100 Meranoplus castaneus 64 Meranoplus radamae 100 Pristomyrmex orbiceps Pristomyrmex punctatus Paratopula TH01 98 Poecilomyrma myrmecodiae Romblonella scrobifera 100 Rotastruma recava Rotastruma stenoceps 96 Diplomorium longipenne 98 Carebara vidua 74 Carebara nosindambo 100 Carebara alperti 91 Carebara AU01 100 Carebara urichi 77 Carebara affinis 100 Myrmecina PG01 100 Myrmecina americana 57 Myrmecina graminicola 97 Crematogaster acuta 100 Crematogaster sordidula 100 Crematogaster modiglianii 100 Crematogaster adrepens Crematogaster emeryana Atopomyrmex mocquerysi 99 100 Terataner bottegoi Terataner MG02 89 Terataner MG27 100 Cataulacus ebrardi 70 Cataulacus hispidulus 100 Ocymyrmex fortior cf 100 Cardiocondyla mauritanica 99 Cardiocondyla MY01 Cardiocondyla thoracica Acanthomyrmex ferox 63 Podomyrma silvicola Xenomyrmex floridanus 58 Huberia brounii 100 Liomyrmex gestroi 53 100 Metapone madagascarica Metapone PG01 100 Eutetramorium mocquerysi 100 Royidris notorthotenes 100 Royidris longiseta 100 Malagidris jugum 100 Malagidris belti Myrmisaraka brevis Vitsika suspicax 54 Recurvidris TH01 74 Vombisidris bilongrudi 100 Nesomyrmex echinatinodis 100 Nesomyrmex wilda 79 100 Nesomyrmex madecassus 55 Nesomyrmex MG01 Nesomyrmex MG08 Gauromyrmex acanthinus 100 Harpagoxenus sublaevis 79 89 Formicoxenus diversipilosus Leptothorax muscorum complex 69 Chalepoxenus kutteri 100 Myrmoxenus bernardi 65 Temnothorax poeyi Temnothorax rugatulus Figure S2 UBx

Apterogyna ZA01 100 Myrmicine genus26 PH02 Vollenhovia emeryi Calyptomyrmex kaurus Melissotarsus insularis 100 Cardiocondyla mauritanica 96 Cardiocondyla MY01 Cardiocondyla thoracica 88 Tetramorium CF01 Tetramorium taylori 89 100 Rhoptromyrmex transversinodis Tetramorium MG125 99 Anergates atratulus 84 100 Teleutomyrmex kutteri 94 Strongylognathus testaceus 96 Tetramorium caespitum cf Tetramorium venator 86 Tetramorium validiusculum 88 100 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 Xenomyrmex floridanus 100 Cataulacus ebrardi Cataulacus hispidulus 100 Dilobocondyla borneensis cf Secostruma MY01 100 Pristomyrmex orbiceps Pristomyrmex punctatus 100 Monomorium criniceps 77 Monomorium destructor Monomorium robustior 100 Terataner bottegoi 98 Terataner MG02 Terataner MG27 100 Ancyridris polyrhachioides nr 66 Lordomyrma bhutanensis cf Lordomyrma desupra Propodilobus pingorum 71 Cyphoidris exalta 100 Myrmecina PG01 96 Myrmecina americana Myrmecina graminicola 58 Lophomyrmex ambiguus Ocymyrmex fortior cf 100 Meranoplus castaneus Meranoplus radamae 100 Poecilomyrma myrmecodiae 100 Romblonella scrobifera 83 Rotastruma recava Rotastruma stenoceps 96 Crematogaster acuta 98 Crematogaster sordidula 79 Crematogaster modiglianii 71 Crematogaster adrepens Crematogaster emeryana 97 Huberia brounii Podomyrma silvicola 100 Liomyrmex gestroi 100 Metapone madagascarica Metapone PG01 100 Nesomyrmex echinatinodis 91 Nesomyrmex wilda 100 Nesomyrmex MG01 76 Nesomyrmex madecassus Nesomyrmex MG08 97 Diplomorium longipenne 100 Carebara vidua 57 Carebara nosindambo 100 Carebara alperti 55 Carebara AU01 100 Carebara urichi Carebara affinis 61 Eutetramorium mocquerysi Malagidris jugum 69 Myalagidris belti Myrmisaraka brevis 100 Vitsika suspicax Royidris notorthotenes Royidris longiseta 59 Vombisidris bilongrudi 99 Formicoxenus diversipilosus 97 Harpagoxenus sublaevis Leptothorax muscorum complex 90 Gauromyrmex acanthinus 100 Myrmoxenus bernardi 78 Chalepoxenus kutteri Temnothorax poeyi Temnothorax rugatulus 100 Stenamma dyscheres Stenamma expolitum 66 Stenamma striatulum 100 Aphaenogaster albisetosa 65 Messor andrei 100 Aphaenogaster pythia 95 Aphaenogaster swammerdami Aphaenogaster araneoides 64 100 Goniomma blanci Oxyopomyrmex santschii 71 Aphaenogaster occidentalis 100 Messor denticornis 87 Messor wasmanni 100 Pogonomyrmex angustus 68 Hylomyrma blandiens Pogonomyrmex imberbiculus 87 Pogonomyrmex striatinodus Pogonomyrmex subdentatus Pogonomyrmex vermiculatus 100 Manica bradleyi 90 100 Manica rubida 62 Myrmica kotokui 63 Myrmica rugiventris 72 Myrmica striolagaster Myrmica tahoensis 99 Acanthoponera minor 90 Rhytidoponera chalybaea 100 Lasius californicus 58 Myrmelachista flavocotea 81 100 Tetraponera rufonigra 79 Aneuretus simoni Dolichoderus pustulatus 78 Myrmecia pyriformis 53 Nothomyrmecia macrops Platythyrea mocquerysi 91 100 100 Ankylomyrma coronacantha 88 Tatuidris tatusia Tatuidris tatusia_2 89 Amblyopone australis Proceratium stictum 97 Paraponera clavata Stigmatomma pallipes Dolopomyrmex pilatus Rogeria nevadensis Stegomyrmex manni Adelomyrmex paratristani Austromorium flavigaster Baracidris sitra Bariamyrma hispidula Megalomyrmex silvestrii 99 Monomorium antarcticum Monomorium brocha Monomorium denticulatum Monomorium hanneli Monomorium latinode Monomorium sechellense Oxyepoecus vezenyii Rogeria stigmatica Tyrannomyrmex legatus 66 91 Adelomyrmex SC01 Cryptomyrmex boltoni 100 Anillomyrma AFRC TZ 01 Anillomyrma decamera 85 Monomorium ferodens Monomorium fisheri 100 Myrmicaria carinata Myrmicaria exigua 100 Monomorium pharaonis Monomorium exiguum 80 61 Monomorium ergatogyna Monomorium madecassum 75 Monomorium fieldi nr 100 Monomorium MG04 Kempfidris inusualis 99 Tropidomyrmex elianae Carebarella PA01 90 Solenopsis papuana 72 Solenopsis fugax 100 Solenopsis ZA02 94 Solenopsis mameti 100 Solenopsis xyloni 100 Solenopsis molesta Solenopsis succinea Acanthognathus ocellatus Apterostigma auriculatum Daceton armigerum Lachnomyrmex scrobiculatus 60 Diaphoromyrma sofiae Lenomyrmex colwelli 71 Mycocepurus goeldii Myrmicocrypta infuscata cf 85 Blepharidatta conops 100 Allomerus octoarticulatus Wasmannia auropunctata 100 Procryptocerus scabriusculus 100 Cephalotes atratus 100 Cephalotes texanus 97 Ochetomyrmex neopolitus 100 Ochetomyrmex semipolitus Tranopelta subterranea Tranopelta gilva 88 Microdaceton tibialis 97 Colobostruma unicorna Orectognathus versicolor 100 Epopostruma monstrosa 54 Mesostruma browni Cyphomyrmex longiscapus cf 78 Kalathomyrmex emeryi Mycetarotes acutus Mycetosoritis hartmanni 58 Cyphomyrmex cornutus 100 Trachymyrmex arizonensis 61 Acromyrmex versicolor Atta texana Pheidole fimbriata 100 Pheidole rhea Machomyrma dispar Pheidole clydei 84 Pheidole longispinosa 70 Anisopheidole antipodum Pheidole hyatti 99 Pheidole lucioccipitalis Pheidole pallidula 72 91 Basiceros manni 100 Protalaridris armata Rhopalothrix isthmica 81 Octostruma EC01 100 Eurhopalothrix australis 100 Eurhopalothrix gravis 80 Talaridris mandibularis 89 Phalacromyrmex fugax Pilotrochus besmerus 83 Strumigenys ambatrix 100 83 Strumigenys nitens Strumigenys gundlachi Strumigenys biolleyi 100 Strumigenys chiricahua Strumigenys exiguaevitae Strumigenys maxillaris Strumigenys membranifera Strumigenys ocypete 51 98 Strumigenys olsoni Strumigenys erynnes Strumigenys ludovici 90 Strumigenys chyzeri 75 Strumigenys emmae Strumigenys godeffroyi 100 Strumigenys rogeri 77 Strumigenys coveri Strumigenys dicomas Figure S2 Wg

Apterogyna ZA01 Amblyopone australis Proceratium stictum Stigmatomma pallipes 99 Platythyrea mocquerysi 91 Paraponera clavata 100 100 Ankylomyrma coronacantha Tatuidris tatusia Tatuidris tatusia_2 100 Lasius californicus Myrmelachista flavocotea 100 Acanthoponera minor Rhytidoponera chalybaea 64 Aneuretus simoni 62 Dolichoderus pustulatus 100 Tetraponera rufonigra 100 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 Xenomyrmex floridanus 81 Adelomyrmex paratristani Baracidris sitra 100 Allomerus octoarticulatus Wasmannia auropunctata 72 Austromorium flavigaster 90 Monomorium antarcticum 58 Carebara urichi Carebara affinis 100 Cataulacus ebrardi Cataulacus hispidulus 60 Lophomyrmex ambiguus Recurvidris TH01 100 Meranoplus castaneus Meranoplus radamae 80 Monomorium brocha Monomorium latinode 54 Monomorium ferodens 100 Monomorium fisheri Royidris notorthotenes Royidris longiseta 100 Myrmicaria carinata 100 Myrmicaria exigua Malagidris jugum Malagidris belti 93 Phalacromyrmex fugax Pilotrochus besmerus 51 Apterostigma auriculatum Microdaceton tibialis Mycocepurus goeldii 90 Carebara nosindambo 100 Carebara alperti Carebara AU01 100 Procryptocerus scabriusculus 100 Cephalotes atratus Cephalotes texanus 74 99 Huberia brounii Myrmisaraka brevis Vitsika suspicax 100 Liomyrmex gestroi 100 Metapone madagascarica Metapone PG01 100 Monomorium robustior 98 Monomorium criniceps Monomorium destructor 100 Myrmecina graminicola 53 Myrmecina americana Myrmecina PG01 100 Terataner bottegoi 74 Terataner MG02 Terataner MG27 Ancyridris polyrhachioides nr 100 Lordomyrma bhutanensis cf Lordomyrma desupra 100 Propodilobus pingorum 91 Anillomyrma AFRC TZ 01 60 Anillomyrma decamera Kempfidris inusualis Tropidomyrmex elianae 67 Kartidris sparsipila 100 Cardiocondyla mauritanica 100 Cardiocondyla MY01 Cardiocondyla thoracica 50 Carebara vidua 100 64 Mesostruma browni 65 Colobostruma unicorna Epopostruma monstrosa Chalepoxenus kutteri 98 Myrmoxenus bernardi Temnothorax poeyi Temnothorax rugatulus 71 Vombisidris bilongrudi 100 Formicoxenus diversipilosus Harpagoxenus sublaevis 100 Leptothorax muscorum complex 100 Ochetomyrmex neopolitus 100 Ochetomyrmex semipolitus Tranopelta subterranea Tranopelta gilva 97 Romblonella scrobifera 94 Poecilomyrma myrmecodiae 100 Rotastruma recava Rotastruma stenoceps 73 Crematogaster acuta 100 Crematogaster sordidula 97 Crematogaster modiglianii 100 Crematogaster adrepens Crematogaster emeryana 100 Dilobocondyla borneensis cf 59 Secostruma MY01 62 Perissomyrmex snyderi 100 Pristomyrmex orbiceps Pristomyrmex punctatus 100 Nesomyrmex echinatinodis 100 Nesomyrmex wilda 100 Nesomyrmex madecassus Nesomyrmex MG01 Nesomyrmex MG08 96 97 Eurhopalothrix australis 91 Eurhopalothrix gravis Talaridris mandibularis 76 Octostruma EC01 100 Protalaridris armata Rhopalothrix isthmica 100 Monomorium pharaonis 88 Monomorium ergatogyna 95 Monomorium fieldi nr 75 Monomorium madecassum 53 Monomorium exiguum Monomorium MG04 69 83 Solenopsis mameti 99 Solenopsis fugax Solenopsis ZA02 71 Solenopsis xyloni 62 Solenopsis molesta Solenopsis succinea Kalathomyrmex emeryi 98 Mycetarotes acutus Mycetosoritis hartmanni 77 Cyphomyrmex cornutus 100 Cyphomyrmex longiscapus cf 100 Acromyrmex versicolor 69 Atta texana Trachymyrmex arizonensis 92 Pheidole fimbriata 100 Pheidole rhea Pheidole longispinosa 100 Anisopheidole antipodum 91 Pheidole hyatti Pheidole lucioccipitalis Pheidole pallidula 73 Machomyrma dispar Pheidole clydei Tetramorium MG125 100 Rhoptromyrmex transversinodis Tetramorium CF01 93 Anergates atratulus 56 Strongylognathus testaceus Teleutomyrmex kutteri Tetramorium caespitum cf 75 Tetramorium taylori 59 Tetramorium spinosum 81 Tetramorium venator 69 Tetramorium severini Tetramorium validiusculum Lenomyrmex colwelli 97 Strumigenys ambatrix 100 88 100 Strumigenys nitens Strumigenys gundlachi Strumigenys biolleyi 99 Strumigenys exiguaevitae Strumigenys maxillaris Strumigenys olsoni 99 95 Strumigenys rogeri Strumigenys erynnes Strumigenys ludovici 99 Strumigenys coveri Strumigenys dicomas 91 Strumigenys ocypete 99 Strumigenys membranifera Strumigenys chiricahua 58 Strumigenys chyzeri Strumigenys emmae Strumigenys godeffroyi 100 Hylomyrma blandiens Pogonomyrmex angustus 86 Pogonomyrmex striatinodus 94 Pogonomyrmex imberbiculus 66 100 Pogonomyrmex subdentatus Pogonomyrmex vermiculatus Manica bradleyi 100 Manica rubida 100 Myrmica kotokui Myrmica rugiventris 57 74 Myrmica striolagaster Myrmica tahoensis 100 Aphaenogaster pythia Aphaenogaster swammerdami 79 Aphaenogaster araneoides 100 Aphaenogaster albisetosa 100 Messor andrei 100 Aphaenogaster occidentalis 97 Messor denticornis Messor wasmanni 100 Goniomma blanci 84 Oxyopomyrmex santschii 97 Stenamma dyscheres Stenamma expolitum Stenamma striatulum Figure S2 EF1aF2

Apterogyna ZA01 Acanthoponera minor Amblyopone australis Paraponera clavata 55 Platythyrea mocquerysi Rhytidoponera chalybaea 83 Aneuretus simoni 91 Dolichoderus pustulatus 100 Lasius californicus 81 Myrmelachista flavocotea 100 Myrmecia pyriformis Nothomyrmecia macrops 53 Proceratium stictum 86 Stigmatomma pallipes 54 Tetraponera rufonigra 94 100 Ankylomyrma coronacantha Tatuidris tatusia Tatuidris tatusia_2 100 Pogonomyrmex angustus 54 Hylomyrma blandiens 81 Pogonomyrmex striatinodus 91 78 Pogonomyrmex subdentatus 84 Pogonomyrmex imberbiculus Pogonomyrmex vermiculatus 99 Manica bradleyi 99 Manica rubida Myrmica kotokui 100 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 Xenomyrmex floridanus 54 Calyptomyrmex kaurus Cyphoidris exalta 100 Cataulacus ebrardi Cataulacus hispidulus 97 Cephalotes atratus Cephalotes texanus 100 Dicroaspis KM01 Proatta butteli 65 Melissotarsus insularis Rhopalomastix rothneyi 55 Myrmicine genus26 PH02 Vollenhovia emeryi 99 Phalacromyrmex fugax Pilotrochus besmerus 100 Pristomyrmex orbiceps Pristomyrmex punctatus 52 Perissomyrmex snyderi 100 Dilobocondyla borneensis cf 76 Secostruma MY01 98 Eurhopalothrix australis 100 Eurhopalothrix gravis Talaridris mandibularis 100 Harpagoxenus sublaevis 68 Formicoxenus diversipilosus Leptothorax muscorum complex 57 Blepharidatta conops Microdaceton tibialis 100 Allomerus octoarticulatus Wasmannia auropunctata 99 Orectognathus versicolor 77 Colobostruma unicorna 54 Epopostruma monstrosa 100 Mesostruma browni 68 Ochetomyrmex neopolitus 100 Ochetomyrmex semipolitus Tranopelta subterranea Tranopelta gilva 96 Romblonella scrobifera 94 Poecilomyrma myrmecodiae 100 Rotastruma recava Rotastruma stenoceps 100 Meranoplus castaneus 65 Meranoplus radamae 100 Cardiocondyla mauritanica 100 Cardiocondyla MY01 Cardiocondyla thoracica 100 Crematogaster sordidula 51 Crematogaster acuta 100 Crematogaster modiglianii 100 Crematogaster adrepens Crematogaster emeryana 100 Nesomyrmex echinatinodis 97 Nesomyrmex wilda 98 Nesomyrmex madecassus 88 Nesomyrmex MG01 Nesomyrmex MG08 Lophomyrmex ambiguus 75 79 Adlerzia froggatti Recurvidris TH01 100 Terataner bottegoi 91 Terataner MG02 Terataner MG27 99 Carebara vidua 99 Carebara alperti 100 Carebara AU01 54 Carebara nosindambo 100 Carebara urichi Carebara affinis Kalathomyrmex emeryi 78 Mycetarotes acutus 77 Mycetosoritis hartmanni 99 Cyphomyrmex longiscapus cf 76 Cyphomyrmex cornutus 100 Trachymyrmex arizonensis 79 Acromyrmex versicolor Atta texana Gauromyrmex acanthinus 72 100 100 Myrmecina PG01 92 Myrmecina americana Myrmecina graminicola 53 Temnothorax poeyi 76 Temnothorax rugatulus 99 Chalepoxenus kutteri Myrmoxenus bernardi 99 Pheidole fimbriata Pheidole rhea 99 86 Anisopheidole antipodum 100 Pheidole hyatti Machomyrma dispar 52 Pheidole clydei Pheidole longispinosa 60 Pheidole lucioccipitalis Pheidole pallidula 99 Stenamma expolitum 98 Stenamma dyscheres 80 Stenamma striatulum 100 Aphaenogaster albisetosa Messor andrei 58 100 Aphaenogaster pythia Aphaenogaster swammerdami 93 60 Aphaenogaster araneoides 100 Goniomma blanci Oxyopomyrmex santschii 99 Aphaenogaster occidentalis 100 Messor denticornis 100 Messor wasmanni 69 Malagidris jugum Malagidris belti 51 100 Eutetramorium mocquerysi Myrmisaraka brevis Vitsika suspicax 97 Huberia brounii 69 Podomyrma silvicola 100 Liomyrmex gestroi 100 Metapone madagascarica 100 Metapone PG01 73 Royidris notorthotenes Royidris longiseta 100 Monomorium robustior 97 Monomorium criniceps Monomorium destructor Acanthomyrmex ferox Mayriella ebbei 100 Ancyridris polyrhachioides nr 80 Lordomyrma bhutanensis cf 60 Lordomyrma desupra 100 Propodilobus pingorum Tetramorium venator Tetramorium validiusculum 100 Tetramorium CF01 Tetramorium taylori 100 100 Tetramorium caespitum cf 78 Strongylognathus testaceus 92 Anergates atratulus Teleutomyrmex kutteri 69 Rhoptromyrmex transversinodis Tetramorium MG125 51 Tetramorium severini Tetramorium spinosum Strumigenys ambatrix 97 98 Strumigenys nitens Strumigenys gundlachi 96 100 Strumigenys biolleyi Strumigenys erynnes Strumigenys ludovici 100 100 Strumigenys rogeri 64 Strumigenys coveri Strumigenys dicomas 96 Strumigenys maxillaris 67 96 Strumigenys olsoni Strumigenys membranifera 54 Strumigenys ocypete 63 Strumigenys chiricahua 70 Strumigenys exiguaevitae 99 Strumigenys godeffroyi 98 Strumigenys chyzeri Strumigenys emmae Rogeria nevadensis 90 Bariamyrma hispidula Rogeria stigmatica 100 Carebarella PA01 99 Solenopsis papuana Solenopsis mameti 100 100 Solenopsis fugax Solenopsis ZA02 97 Solenopsis molesta 62 73 Solenopsis succinea Solenopsis xyloni Megalomyrmex silvestrii Monomorium brocha Monomorium hanneli Tyrannomyrmex legatus 100 Anillomyrma AFRC TZ 01 Anillomyrma decamera 100 Myrmicaria carinata Myrmicaria exigua 64 88 Monomorium denticulatum 53 Austromorium flavigaster 76 Monomorium antarcticum 63 Adelomyrmex paratristani 100 Oxyepoecus vezenyii Kempfidris inusualis Tropidomyrmex elianae Baracidris sitra Monomorium sechellense 57 51 Adelomyrmex SC01 Cryptomyrmex boltoni 75 Monomorium ferodens 87 Monomorium fisheri 77 Monomorium latinode Monomorium madecassum 85 Monomorium MG04 Monomorium pharaonis 69 Monomorium ergatogyna 60 Monomorium exiguum Monomorium fieldi nr Figure S2 CAD

Apterogyna ZA01 Platythyrea mocquerysi 100 Proceratium stictum Stigmatomma pallipes 100 100 Ankylomyrma coronacantha 65 Tatuidris tatusia Tatuidris tatusia_2 93 Amblyopone australis 65 Paraponera clavata Tetraponera rufonigra 69 Acanthoponera minor 57 Rhytidoponera chalybaea 55 87 Aneuretus simoni Dolichoderus pustulatus 100 Lasius californicus 94 Myrmelachista flavocotea 100 Myrmecia pyriformis 65 Nothomyrmecia macrops 100 Pogonomyrmex vermiculatus 68 Hylomyrma blandiens 100 Pogonomyrmex angustus 100 Pogonomyrmex striatinodus 100 100 Pogonomyrmex imberbiculus Pogonomyrmex subdentatus 100 Manica bradleyi 100 Manica rubida 100 Myrmica kotokui 67 96 Myrmica rugiventris 91 Myrmica striolagaster Myrmica tahoensis 100 Aphaenogaster albisetosa Messor andrei 100 100 Stenamma dyscheres Stenamma expolitum Stenamma striatulum 98 98 Aphaenogaster araneoides 100 Aphaenogaster occidentalis 93 100 Messor denticornis Messor wasmanni 100 Aphaenogaster pythia 91 Aphaenogaster swammerdami 100 Goniomma blanci Oxyopomyrmex santschii 92 Stegomyrmex manni 70 Dolopomyrmex pilatus Rogeria stigmatica 72 Bariamyrma hispidula 100 Rogeria nevadensis 100 Monomorium denticulatum Oxyepoecus vezenyii 93 95 Megalomyrmex silvestrii 66 Austromorium flavigaster 100 Monomorium antarcticum 87 Monomorium brocha 76 Monomorium latinode 100 Tyrannomyrmex legatus Kempfidris inusualis 94 Tropidomyrmex elianae 100 Carebarella PA01 98 Solenopsis papuana 100 Solenopsis xyloni 100 100 Solenopsis fugax Solenopsis ZA02 61 Solenopsis mameti 100 Solenopsis molesta 92 Solenopsis succinea Monomorium ferodens Monomorium fisheri Monomorium hanneli 100 Myrmicaria carinata Myrmicaria exigua 66 Monomorium sechellense 100 100 Anillomyrma AFRC TZ 01 Anillomyrma decamera 99 Adelomyrmex paratristani Baracidris sitra 57 Adelomyrmex SC01 Cryptomyrmex boltoni 100 Monomorium pharaonis 57 Monomorium madecassum 78 Monomorium ergatogyna 89 Monomorium fieldi nr 100 Monomorium exiguum Monomorium MG04 100 Orectognathus versicolor 100 Colobostruma unicorna 77 100 Epopostruma monstrosa 99 Mesostruma browni 99 Ochetomyrmex neopolitus 100 Ochetomyrmex semipolitus Tranopelta subterranea 58 Tranopelta gilva Acanthognathus ocellatus Diaphoromyrma sofiae Lachnomyrmex scrobiculatus 72 Lenomyrmex colwelli Microdaceton tibialis Apterostigma auriculatum 91 Daceton armigerum 74 99 Mycocepurus goeldii Myrmicocrypta infuscata cf 82 Kalathomyrmex emeryi 61 Mycetarotes acutus 100 Mycetosoritis hartmanni 100 Cyphomyrmex cornutus 98 63 Cyphomyrmex longiscapus cf 100 Trachymyrmex arizonensis 84 Acromyrmex versicolor Atta texana 100 Blepharidatta conops 100 Allomerus octoarticulatus Wasmannia auropunctata 100 100 Procryptocerus scabriusculus 92 Cephalotes atratus Cephalotes texanus 100 Pheidole fimbriata 97 Pheidole rhea 100 Pheidole clydei 100 69 Anisopheidole antipodum 80 66 Machomyrma dispar Pheidole longispinosa 80 Pheidole hyatti 57 Pheidole lucioccipitalis Pheidole pallidula Basiceros manni 99 100 Protalaridris armata Rhopalothrix isthmica 99 Octostruma EC01 99 Eurhopalothrix australis 99 100 Eurhopalothrix gravis Talaridris mandibularis 100 Phalacromyrmex fugax 98 Pilotrochus besmerus Strumigenys ambatrix 100 100 100 Strumigenys nitens Strumigenys gundlachi Strumigenys biolleyi 99 Strumigenys exiguaevitae Strumigenys maxillaris 100 Strumigenys olsoni Strumigenys erynnes 100 Strumigenys ludovici 100 Strumigenys rogeri 99 Strumigenys coveri Strumigenys dicomas 100 100 Strumigenys chiricahua 77 Strumigenys membranifera Strumigenys ocypete 99 Strumigenys godeffroyi 100 Strumigenys chyzeri 59 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 Tetheamyrma subspongia 100 Cataulacus ebrardi Cataulacus hispidulus 53 Dacatria templaris Diplomorium longipenne 100 Dilobocondyla borneensis cf Secostruma MY01 100 Melissotarsus insularis Rhopalomastix rothneyi 100 Meranoplus castaneus Meranoplus radamae 99 Pristomyrmex orbiceps Pristomyrmex punctatus 64 Adlerzia froggatti 100 Myrmicine genus26 PH02 Vollenhovia emeryi 91 Proatta butteli Vombisidris bilongrudi Xenomyrmex floridanus 100 Terataner bottegoi 54 Terataner MG02 Terataner MG27 100 Ancyridris polyrhachioides nr 69 Propodilobus pingorum 73 Lordomyrma bhutanensis cf Lordomyrma desupra 100 Crematogaster acuta 100 Crematogaster sordidula 99 Crematogaster modiglianii 100 Crematogaster adrepens 95 Crematogaster emeryana 100 Nesomyrmex echinatinodis 100 Nesomyrmex wilda 100 Nesomyrmex MG08 88 Nesomyrmex madecassus Nesomyrmex MG01 63 Paratopula TH01 100 Poecilomyrma myrmecodiae 69 Romblonella scrobifera 100 Rotastruma recava Rotastruma stenoceps 100 59 Cardiocondyla mauritanica 67 Cardiocondyla MY01 Cardiocondyla thoracica 100 Myrmecina PG01 100 Myrmecina americana Myrmecina graminicola 96 Carebara vidua 97 Carebara nosindambo 100 Carebara alperti 52 Carebara AU01 100 Carebara urichi Carebara affinis 100 76 Leptothorax muscorum complex 100 Formicoxenus diversipilosus Harpagoxenus sublaevis 100 Temnothorax poeyi 81 Temnothorax rugatulus 99 Chalepoxenus kutteri Myrmoxenus bernardi 100 Tetramorium CF01 Tetramorium taylori 100 86 Rhoptromyrmex transversinodis 100 100 Tetramorium MG125 82 Tetramorium severini 100 Tetramorium spinosum Tetramorium venator 73 Tetramorium validiusculum 100 Tetramorium caespitum cf 100 Strongylognathus testaceus 100 Anergates atratulus Teleutomyrmex kutteri 100 Monomorium robustior 100 Monomorium criniceps Monomorium destructor 100 100 Eutetramorium mocquerysi Royidris notorthotenes 100 Royidris longiseta 97 80 Malagidris jugum 100 Malagidris belti Myrmisaraka brevis Vitsika suspicax 100 Podomyrma silvicola 58 Huberia brounii 100 Liomyrmex gestroi 100 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

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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

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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

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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

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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

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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

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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

1References 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.

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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

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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-

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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

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