The evolution of social in revealed by a global phylogeny – Supplementary figures, tables, and references

Marek L. Borowiec Stefan P. Cover Christian Rabeling

1 Supplementary Methods

Data availability Trimmed reads generated for this study are available at the NCBI Sequence Read Archive (to be submit­ ted upon publication). Detailed voucher collection information, assembled sequences, analyzed matrices, configuration files and output of all analyses, and code used are available on Zenodo (DOI: 10.5281/zen­ odo.4341310).

Taxon sampling For this study we gathered samples collected in the past ~60 years which were available as either ethanol­ preserved or point­mounted specimens. Taxon sampling comprises 101 newly sequenced ingroup morphos­ pecies from all seven groups of Formica ants Creighton (1950) that were recognized prior to our study and 8 outgroup species. Our sampling was guided by previous taxonomic and phylogenetic work Creighton (1950); Francoeur (1973); Snelling and Buren (1985); Seifert (2000, 2002, 2004); Goropashnaya et al. (2004, 2012); Trager et al. (2007); Trager (2013); Seifert and Schultz (2009a,b); Muñoz­López et al. (2012); Antonov and Bukin (2016); Chen and Zhou (2017); Romiguier et al. (2018) and included represen­ tatives from both the New and the Old World. Collection data associated with sequenced samples can be found in Table S1.

Molecular data collection and sequencing We performed non­destructive extraction and preserved same­specimen vouchers for each newly sequenced sample. We re­mounted all vouchers, assigned unique specimen identifiers (Table S1), and deposited them in the ASU Social Repository (contact: Christian Rabeling, [email protected]). Detailed voucher collection data can be accessed at 10.5281/zenodo.4341310. Briefly, we extracted DNA from all newly sequenced specimens using a DNeasy Blood and Tissue Kit (Qiagen, Valencia, CA, USA) and followed a library preparation protocol that follows was slightly modified from Blaimer et al. (2015). We used a KAPA Hyper Prep Library Kit (Kapa Biosystems, Inc., Wilmington, MA, USA) with magnetic bead cleanup Fisher et al. (2011) and a SPRI substitute Rohland and Reich (2012) as described in Faircloth et al. (2014). We used TruSeq adapters Faircloth and Glenn (2012) for ligation followed by PCR amplification of the library using a mix of HiFi HotStart polymerase reaction mix (Kapa Biosystems), Illumina TruSeq primers, and nuclease­free water. We enriched each pool with 9,446 custom­designed probes (MYcroarray, Inc.) targeting 2,524 UCE loci in Branstetter et al. (2017). We followed library enrichment procedures for the MYcroarray MYBaits kit Blumenstiel et al. (2010) except we used a 0.1× of the standard MYBaits concentration and added 0.7 µL of 500 µM custom blocking oligos designed against the custom sequence tags. We ran the hybridization reaction for 24 h at 65 °C, subsequently bound all pools to streptavidin beads (MyOne C1; Life Technologies) and washed bound libraries according to a standard target enrichment protocol Blumenstiel et al. (2010). We used the with­bead approach for PCR recovery of enriched libraries as described in Faircloth (2015). We submitted pre­pooled libraries to the University of Utah High Throughput Genomics Core Facility for quality control on Agilent Bioanalyzer (Agilent Technologies, Santa Clara, CA, USA) and normalization. The pooled libraries were then sequenced using one full and one partial lane of a HiSeq 125 Cycle Paired­ End Sequencing v4 run. Quality­trimmed sequence reads generated as part of this study are available from the NCBI Sequence Read Archive (to be submitted upon acceptance for publication).

2 Processing of UCE data We performed read cleaning, assembly, and matching of assembled contigs to UCE probes using Phyluce bioinformatics pipeline Faircloth (2015). We trimmed the FASTQ data using Illumiprocessor, a wrapper around Trimmomatic Bolger et al. (2014), with default settings (LEADING:5, TRAILING:15, SLIDING­ WINDOW:4:15, MINLEN:40). Assemblies were done using Trinity v20140717 Grabherr et al. (2011) with the phyluce_assembly_assemblo_trinity wrapper. We then assessed orthology by matching the assembled contigs to enrichment probe sequences with phyluce_assembly_match_contigs_to_probes (min_coverage=50, min_identity=80).

Alignment We used phylogeny­aware UPP workflow Nguyen et al. (2015) to align all UCE sequences. We used AMAS Borowiec (2016) for alignment wrangling and obtaining summary statistics and AliView Larsson (2014) for visualization. Although alignment trimming has been criticized in the past Tan et al. (2015), we decided to trim the alignments because of substantial computational burden associated with analysis of untreated data with high proportion of gaps. We used trimAl Capella­Gutierrez et al. (2009) and its “gappyout” algo­ rithm, which is a relatively relaxed algorithm for removal of gappy sites. Visual inspection of alignments revealed that occasionally sequences were misaligned towards flanks. To automatically identify and discard the misaligned sequences we wrote a custom R script (R Core Team) that leveraged packages ape Paradis et al. (2004); Paradis (2012), seqinr Charif and Lobry (2007), doParallel, and plyr. The script first generates a matrix of uncorrected p­distances from a UCE locus alignment and for each taxon it computes average p­distance to all other taxa. Then it creates a distribution of average per­locus p­distances for each taxon and detects outliers defined as sequences that lay above 3 SD from the mean of that distribution. Once identified, the script removes outliers using AMAS. This procedure resulted in removal of 0.97 % of all sequences. For downstream analyses we retained only alignments that had 110 or more taxa (70 % of total), resulting in 2,242 loci on average 667 nt long. The resulting concatenated matrix was 1,497,044 nt long and contained 17.58 % of missing data and gaps.

Partitioning We used ModelFinder Kalyaanamoorthy et al. (2017) as implemented in IQ­TREE Nguyen et al. (2014). For each UCE we selected the best model under AICc. These models were then used for by­locus partitioned analysis of concatenated data matrix Chernomor et al. (2016). We have also employed the newly proposed strategy of partitioning UCE loci based on a sliding window approach that groups UCE sites with simi­ lar entropies Tagliacollo and Lanfear (2018). Unfortunately, many partitions identified using this approach were saturated and caused numerical instability in maximum likelihood analyses using IQ­TREE, result­ ing in unreasonably long tree lengths. We therefore proceeded with downstream analyses using per­locus partitioning.

Phylogenetic and concordance analyses using maximum likelihood We used IQ­TREE Nguyen et al. (2014) for maximum likelihood inference of phylogeny on single­locus alignments and concatenated data matrix. To test the robustness of the partitioned concatenated analysis, we performed unpartitioned analysis under HKY+4G model, which was the most common model identified as best under AICc for single loci. To assess the sensitivity of our results using measures other than bootstrap support we performed a quartet sampling analysis with 500 maximum replicates (Pease et al., 2018). Results are summarized in Figure S7.

3 Species tree analyses In addition to concatenated analyses we performed coalescent­based species tree estimation using ASTRAL­ III (Zhang et al., 2018). Because summary coalescence methods such as ASTRAL have been shown to be negatively impacted by error in estimated gene trees Roch and Warnow (2015) we used the weighted sta­ tistical binning pipeline Mirarab et al. (2014); Bayzid et al. (2015). We collapsed all nodes with ultrafast bootstrap Minh et al. (2013) support below 95 for the binning pipeline, which resulted in identification of 1,733 supergenes containing from one to three UCE loci. We then used IQ­TREE to estimate supergene trees under a fully partitioned model (i.e. with branch lengths unlinked across partitions). Because of re­ cent criticism of the statistical binning pipeline Adams and Castoe (2019) we also performed an analysis where raw trees from individual locus analyses were used. We mapped all terminals to putative species using morphology and the concatenated tree as guidance. Because some of the 101 species we recognized using morphology were non­monophyletic on the concatenated tree, we mapped the terminals onto 113 to­ tal monophyletic lineages representing putative species (Figures S4–S6). To test the effect of missing data Sayyari et al. (2017) on the position of Formica talbotae we performed additional analysis that used only the 67 loci which contained at least 50 % complete sequence for this taxon (Figure S6).

Divergence time inference For divergence time analyses we used a node dating approach, as implemented in MCMCTree, a part of the PAML package, v4.9e Yang (2007). MCMCTree utilized rapid approximate likelihood computation dos Reis and Yang (2011), which makes it suitable for divergence dating of genome­scale data sets dos Reis et al. (2012). We constrained our root node with soft bounds around a conservative maximum age estimate of 79 Ma, which corresponds to the lower bound of the 95 % highest posterior density interval for that split in Blaimer et al. (2015). Although Formica has a rich fossil record, the affinity of these fossils is uncertain because, as this study shows, morphology has thus far been misleading about phylogeny. Because of this, we conservatively constrained the split of and Formica+ to be at least 34 Ma, or one of the younger estimates for the age of Baltic amber Aleksandrova and Zaporozhets (2008a,b). We ran each analysis unpartitioned, under the HKY+4G model for 20 million generations. We examined each run’s statistics in Tracer.

Biogeographic analyses For biogeographic inference we used BioGeoBEARS v1.1.2 Matzke (2013). We discretized the distribution of Formica species into two regions, the New World and Old World. Model selection implemented in Bio­ GeoBEARS suggested DEC+J Matzke (2014) as the best­fitting model and we used it for all downstream analyses. We used 100 replicates of biogeographical stochastic maps Dupin et al. (2017) to estimate the number of times Formica dispersed between the Old and New World. We used our time­calibrated tree with taxon duplicates removed such that each species or putative species was represented by only one terminal. Results are summarized in Figure S2.

Ancestral state reconstruction In order to investigate the evolution of natural history traits in Formica and Polyergus we used stochastic character mapping (Huelsenbeck et al., 2003). We used the same time­calibrated tree as for biogeographic analyses but with distant ( minuchae and mongolica) outgroups and Formica taxa for which we had insufficient life history data (see Table S2) pruned. For each species we collected literature and previously unpublished field research data on nest struc­ ture, colony structure, and colony foundation mode. We discretized nest structure into three categories: i)

4 monodomous, ii) polydomous, and iii) supercolonial. For the analysis we assigned the highest nest structure complexity recorded, meaning that if a species is known to form monodomous, polydomous, or supercolonial nests, we characterized it as supercolonial. We discretized colony structure into i) monogynous or ii) polyg­ ynous. We assigned species to the polygynous category if they have been observed to be either monogynous or polygynous. We discretized colony founding mode into five categories: i) independent colony foundation via haplometrosis or pleometrosis, queen readoption occurs, budding absent or at low frequency, ii) faculta­ tive temporary social parasitism with budding at low frequency, iii) facultative temporary social parasitism with budding at high frequency, iv) obligate temporary social parasitism and dulosis without budding, v) per­ manent social parasitism. The coding of character states for each species and references to original research articles are available in Table S2. We performed ancestral state reconstruction on each of the three characters (nest structure, colony struc­ ture, and colony founding). We first compared three commonly­used variants of the discrete character evolu­ tion Mk model (Lewis, 2001): all rates equal (character state change rates are equal for all states), symmetric transition rates (character state change rates are different for each pair of states), and all rates different (char­ acter state change rates are different for each transition). We also fit the meristic model (assuming characters change in step­wise fashion). We fit these four models using ”fit” functions the R package GEIGER v2.0.6 (Harmon et al., 2007) to all three characters. We used Akaike Information Criterion corrected for sample size (AICc) weights as computed by the ”aic.w” function in Phytools v0.6­44 (Revell, 2011) to see which model fit best. This approach identified the all rates different model to be the best fit for nest and colony structure and the symmetric model was found best­fitting for colony founding. Ancestral reconstruction results are found in Figures S3­S5.

5 Formica_querquetulana_F0081 9 5 100Formica_querquetulana_F0095 7 2 Formica_nepticula_F0077 8 9 Formica_knighti_F0015 Formica_talbotae_F0156 8 9 Formica_densiventris_F0072 100 8 4 Formica_cf_densiventris_F0164 6 9 Formica_densiventris_F0073 Formica_sp_micro_grp_F0087 100 Formica_sp_micro_grp_F0096 Formica_mlb04_micro_grp_F0085 9 8 100 100 Formica_sp_micro_grp_F0091 100 Formica_mlb03_micro_grp_F0084 Formica_mlb02_micro_grp_F0083 Formica_nevadensis_F0079 9 9 100 100 Formica_sp_micro_grp_F0093 109 90 Formica_nevadensis_F0078 Formica_nevadensis_F0080 100 Formica_cf_densiventris_F0075 100 Formica_mlb01_micro_grp_F0082 Formica_adamsi_alpina_F0068 100 10Fo0 rmica_sp_micro_grp_F0089 100 10Fo0rmica_sp_micro_grp_F0090 Formica_sp_micro_grp_F0074 100 100 Formica_sp_micro_grp_F0092 Formica_sp_micro_grp_F0094 Formica_sp_micro_grp_F0157 Formica_obscuripes_F0009 100 100 Formica_obscuripes_F0162 100 Formica_obscuripes_F0008 Formica_obscuripes_F0007 100 100 Formica_subnitens_F0022 100 100 Formica_subnitens_F0025 Formica_cf_subnitens_F0014 Formica_integroides_F0005 8 0 100 100 Formica_propinqua_F0016 100 Formica_ravida_F0027 9 7 Formica_integroides_F0114 100 Formica_ravida_F0026 Formica_coloradensis_F0002 100 Formica_oreas_F0012 100 100Formica_oreas_F0125 100 100 Formica_oreas_F0013 100 Formica_ciliata_F0001 Formica_cf_planipilis_F0028 Formica_fossaceps_F0003 7 8 9 6 Formica_obscuriventris_F0006 9 6 Formica_cf_fossaceps_F0004 100 100 Formica_cf_interga_F0010 Formica_obscuriventris_F0011 Formica_polyctena_F0019 9 9 100 100 Formica_rufa_F0018 100 Formica_lugubris_F0021 100 Formica_pratensis_F0020 Formica_cf_frontalis_F0017 100 Formica_truncorum_F0023 Formica_dakotensis_F0070 100 100 Formica_reflexa_F0086 100 Formica_dakotensis_F0069 Formica_dakotensis_F0071 Formica_sp_sang_grp_F0123 100 Formica_rubicunda_F0054 100 100 Formica_puberula_F0056 100 100 Formica_wheeleri_F0057 Formica_emeryi_F0050 100 Formica_nr_pergandei_F0051 100 100 Formica_puberula_F0122 100 Formica_pergandei_F0053 Formica_rubicunda_F0055 100 Formica_aserva_F0044 100 9 5 Formica_aserva_F0067 100 Formica_aserva_F0163 100 Formica_aserva_F0041 100 9 6 100 Formica_aserva_F0042 Formica_aserva_F0043 Formica_sanguinea_F0048 100 Formica_sanguinea_F0049 Formica_exsectoides_F0032 100 10Formica_exsectoides_F0030 0 100 Formica_exsectoides_F0031 Formica_ulkei_F0029 100 Formica_mesasiatica_F0035 9 0 8 0 100 100 Formica_pressilabris_F0033 Formica_exsecta_F0034 Formica_uralensis_F0024 Formica_neorufibarbis_F0128 9 4 Formica_neorufibarbis_t_F0108 100 Formica_neorufibarbis_F0127 100 100 Formica_neorufibarbis_t_F0126 6 1 Formica_neorufibarbis_F0109 100 100 Formica_neorufibarbis_F0107 Formica_hewitti_F0113 100 100 Formica_hewitti_F0142 Formica_candida_F0155 Formica_clara_F0118 100 100 Formica_rufibarbis_F0136 100 Formica_cunicularia_F0119 Formica_orangea_F0134 Formica_subsericea_F159 8 9 100 Formica_subsericea_F0144 100 Formica_subsericea_F0146 Formica_glacialis_F0121 100 Formica_podzolica_F0145 100 Formica_altipetens_F0099 9 6 100 100 Formica_canadensis_F0102 Formica_cf_altipetens_F0100 10100 0 Formica_neoclara_F0106 Formica_montana_nr_F0117 100 100 Formica_propatula_F0116 7 2 Formica_nr_occulta_F0131 Formica_argentea_F0110 9 6 100 Formica_sp_fusca_grp_F0143 7 9 100 Formica_sp_fusca_grp_F0165 Formica_aerata_F0098 Formica_pacifica_F0135 100 100 Formica_transmontanis_F0141 10100 0 Formica_accreta_F0097 100 Formica_subaenescens_F0104 Formica_xerophila_F0139 100 100 Formica_cf_xerophila_F0140 Formica_cinerea_F0111 100 100 Formica_fuscocinerea_F0115 100 Formica_lemani_F0130 100 Formica_sp_fusca_grp_F0101 Formica_sibylla_F0132 100 Formica_cf_sibylla_F0133 100 Formica_subpolita_F0137 100 Formica_subpolita_F0138 Formica_gnava_F0147 9 7 9 9 Formica_sp_fusca_grp_F0160 9 5 Formica_foreliana_F0103 100 Formica_gnava_F0105 100 Formica_sp_fusca_grp_F0161 Formica_moki_F0129 Formica_dolosa_F0061 100 Formica_dolosa_F0063 100 100 Formica_incerta_F0059 100 100 Formica_incerta_F0065 Formica_pallidefulva_F0060 100 100 Formica_pallidefulva_F0066 100 Formica_archboldi_F0062 Formica_manni_F0037 100 Formica_neogagates_F0038 100 100 100 100 Formica_perpilosa_F0039 Formica_lasioides_F0036 100 Formica_lasioides_F0040 100 Formica_obtusopilosa_F0046 100 100 Formica_obtusopilosa_F0047 100 Formica_sp_neog_grp_F0058 Formica_gagates_F0120 Iberoformica_subrufa_D1135 Polyergus_mexicanus_F0153 100 100 Polyergus_vinosus_F0124 100 Polyergus_mexicanus_F0154 100 Polyergus_longicornis_F0158 Polyergus_rufescens_F0149 Proformica_mongolica_F0150 100 Rossomyrmex_minuchae_F0152

0.007

Supplementary Figure S1: Phylogram of Formica and outgroups from concatenated alignment inferred under max­ imum likelihood using IQ­TREE. Support in ultrafast bootstrap (values above 95 indicate strong support). Scale in substitutions per site.

6 Formica sp micro grp F0093 US-CA6 Formica nevadensis F0079 US-CA3 Formica nevadensis F0078 US-CA2 New World Formica nevadensis F0080 US-CA Formica sp micro grp F0091 US-CA5 Formica mlb04 micro grp F0085 US-CA4 Formica mlb03 micro grp F0084 US-CA3 Old World Formica mlb02 micro grp F0083 US-CA2 Formica mlb01 micro grp F0082 US-CA Formica cf densiventris F0075 US-CA Old World and New World Formica querquetulana F0095 US-MA Formica querquetulana F0081 US-MA Formica nepticula F0077 US-MA Formica knighti F0015 US-MA Formica talbotae F0156 US-ND Formica cf densiventris F0164 US-MT Formica densiventris F0072 US-AZ Formica densiventris F0073 US-CO Formica sp micro grp F0096 US-CO4 Formica sp micro grp F0087 US-CO5 Formica sp micro grp F0089 US-CO3 Formica adamsi alpina F0068 US-CO Formica sp micro grp F0090 US-CO2 Formica sp micro grp F0092 US-UT Formica sp micro grp F0074 US-UT2 Formica sp micro grp F0094 US-CO Formica sp micro grp F0157 US-MI Formica obscuripes F0162 US-MT Formica obscuripes F0009 US-MI Formica obscuripes F0008 US-UT Formica obscuripes F0007 US-WA Formica subnitens F0025 US-CO Formica subnitens F0022 US-CA Formica cf subnitens F0014 US-CA Formica propinqua F0016 US-CA Formica integroides F0005 US-CA2 Formica ravida F0027 US-CA Formica integroides F0114 US-CA Formica ravida F0026 US-CO Formica coloradensis F0002 US-UT Formica cf integra F0010 CA-NS Formica fossaceps cf F0004 CA-NS Formica obscuriventris F0006 US-CO Formica fossaceps F0003 US-UT Formica obscuriventris F0011 US-MI Formica oreas F0125 US-CA Formica oreas F0012 US-CA Formica oreas F0013 US-CO Formica ciliata F0001 US-UT Formica cf planipilis F0028 US-CA F0018 PL F0019 PL F0021 IT Formica pratensis F0020 PL Formica truncorum F0023 PL Formica cf frontalis F0017 KG Formica reflexa F0086 CA-NS Formica dakotensis F0070 US-UT Formica dakotensis F0069 US-AZ Formica dakotensis F0071 US-CA Formica wheeleri F0057 US-AZ Formica puberula F0056 US-UT Formica rubicunda F0054 US-CO Formica sp sang grp F0123 US-MT Formica emeryi F0050 US-CO Formica puberula F0122 US-MT Formica nr pergandei F0051 US-CA F0053 US-MI Formica rubicunda F0055 US-MI Formica aserva F0067 US-UT Formica aserva F0044 US-CO Formica aserva F0163 US-MT Formica aserva F0042 US-WA Formica aserva F0041 US-CA Formica aserva F0043 CA F0049 KG Formica sanguinea F0048 PL Formica exsectoides F0030 US-MA Formica exsectoides F0032 US-NY Formica exsectoides F0031 US-CO Formica ulkei F0029 CA-NS Formica pressilabris F0033 NO Formica mesasiatica F0035 KG Formica exsecta F0034 PL Formica uralensis F0024 PL Formica neorufibarbis t F0126 US-CA2 Formica neorufibarbis F0127 US-CA2 Formica neorufibarbis t F0108 US-CA Formica neorufibarbis t F0128 US-CA3 Formica neorufibarbis F0107 US-CO Formica neorufibarbis F0109 US-CA Formica hewitti F0142 US-CO Formica hewitti F0113 US-CA Formica candida F0155 KG F0136 PL Formica clara F0118 KG F0119 PL Formica orangea F0134 KG Formica subsericea F0144 US-NC Formica subsericea F0159 US-VA Formica subsericea F0146 US-MI Formica podzolica F0145 US-NY F0121 CA-NS Formica canadensis F0102 US-CO Formica altipetens F0099 US-AZ Formica cf altipetens F0100 US-CA Formica neoclara F0106 US-CA Formica propatula F0116 MX-OAX Formica nr montana F0117 MX-OAX Formica nr occulta F0131 US-AZ Formica sp fusca grp F0143 US-CA2 Formica argentea F0110 MX-BCN Formica sp fusca grp F0165 US-MT Formica aerata F0098 US-CA F0104 US-CA Formica accreta F0097 US-CA Formica transmontanis F0141 US-CA Formica pacifica F0135 US-WA Formica cf xerophila F0140 US-CA Formica xerophila F0139 US-AZ Formica fuscocinerea F0115 DE Formica cinerea F0111 PL Formica lemani F0130 PL Formica sp fusca grp F0101 US-CA Formica subpolita F0138 US-UT Formica subpolita F0137 US-CA Formica sibylla cf F0133 US-CA Formica sibylla F0132 US-CA Formica sp fusca grp F0160 US-AZ Formica gnava F0147 US-AZ Formica foreliana F0103 USAZ Formica gnava F0105 US-AZ Formica sp fusca grp F0161 US-NM Formica moki F0129 US-CA F0065 US-MA Formica incerta F0059 US-MI Formica dolosa F0063 US-MA Formica dolosa F0061 US-FL F0066 US-TX Formica pallidefulva F0060 US-MI Formica archboldi F0062 US-FL Formica neogagates F0038 US-CA Formica manni F0037 US-NV Formica perpilosa F0039 US-NM Formica lasioides F0040 US-CA Formica lasioides F0036 US-CA Formica obtusopilosa F0047 US-CO Formica obtusopilosa F0046 US-UT Formica sp neog grp F0058 US-CO Formica gagates F0120 CZ Iberoformica subrufa D1135 ES Polyergus vinosus F0124 US-CA Polyergus mexicanus F0153 US-CA Polyergus mexicanus F0154 US-AZ Polyergus longicornis F0158 US-FL F0149 KG

30 25 20 15 10 5 0 Millions of years ago

Supplementary Figure S2: Biogeographic history of Formica, Iberoformica and Polyergus. Pie charts depict relative likelihoods of range estimations from BioGeoBEARS under DEC+J. Pie charts at nodes correspond to ancestral state estimations and charts on the corners correspond to ranges immediately following speciation. Numbers preceded by the letter F indicate extraction code and correspond to identifiers in Table S1. Country and state/province codes as in main text Figure 1.

7 ● Polyergus rufescens F0149 ● Polyergus longicornis F0158 ● Polyergus vinosus F0124 ● ● Polyergus mexicanus F0153 ● Iberoformica subrufa D1135 ● Formica gagates F0120 ● Formica sp neog grp F0058 ● Formica obtusopilosa F0047 ● Formica lasioides F0040 ● Formica perpilosa F0039 ● Formica neogagates F0038 ● Formica manni F0037 ● ● Formica archboldi F0062 ● ● Formica pallidefulva F0066 ● ● Formica incerta F0059 ● ● Formica dolosa F0061 ● Formica moki F0129 ● Formica gnava F0105 ● ● Formica foreliana F0103 ● ● Formica gnava F0147 ● Formica subpolita F0137 ● Formica lemani F0130 ● ● Formica fuscocinerea F0115 ● Formica cinerea F0111 ● Formica cf xerophila F0140 ● Formica xerophila F0139 ● ● Formica subaenescens F0104 ● ● ● Formica accreta F0097 ● ● Formica transmontanis F0141 ● Formica aerata F0098 ● ● ● Formica argentea F0110 ● Formica nr occulta F0131 ● ● nr F0117 ● Formica neoclara F0106 ● ● ● Formica cf altipetens F0100 ● ● ● Formica canadensis F0102 ● ● ● Formica altipetens F0099 ● ● Formica podzolica F0145 ● ● Formica glacialis F0121 ● Formica subsericea F0144 ● Formica cunicularia F0119 ● Formica rufibarbis F0136 ● Formica clara F0118 ● Formica candida F0155 ● ● Formica hewitti F0113 ● Formica neorufibarbis F0109 ● Formica neorufibarbis t F0126 ●● Formica neorufibarbis F0127 ● Formica neorufibarbis t F0108 ● ● Formica neorufibarbis F0128 ● Formica uralensis F0024 ● Formica exsecta F0034 ● Formica pressilabris F0033 ● Formica ulkei F0029 ● ● Formica exsectoides F0031 ● ● Formica exsectoides F0030 ● Formica sanguinea F0048 ● Formica aserva F0041 ● ● Formica rubicunda F0055 ● Formica pergandei F0053 ● Formica puberula F0122 ● Formica nr pergandei F0051 ● Formica emeryi F0050 ● Formica wheeleri F0057 ● ● ● Formica puberula F0056 ● ● Formica rubicunda F0054 ● Formica dakotensis F0069 ● ● Formica reflexa F0086 ● Formica truncorum F0023 ● Formica pratensis F0020 ● ● Formica lugubris F0021 ● ● Formica rufa F0018 ● ● ● Formica polyctena F0019 ● Formica obscuriventris F0011 ● ● Formica integra F0010 ● ● Formica cf fossaceps F0004 ● ● Formica obscuriventris F0006 ● ● ● Formica fossaceps F0003 ● Formica cf planipilis F0028 ● Formica ciliata F0001 ● Formica oreas F0013 ● ● Formica coloradensis F0002 ● ● Formica ravida F0026 ● ● Formica integroides F0114 ● ● Formica ravida F0027 ● ● ● Formica propinqua F0016 ● ● Formica integroides F0005 ● ● Formica cf subnitens F0014 ● Formica subnitens F0025 ● Formica obscuripes F0007 ● Formica sp micro grp F0157 ● Formica sp micro grp F0094 ● Formica sp micro grp F0092 ●● Formica sp micro grp F0074 ● ● Formica sp micro grp F0090 ●● Formica sp micro grp F0089 ●● Formica adamsi alpina F0068 ● Formica mlb01 micro grp F0082 ● ● Formica cf densiventris F0075 ● ● Formica nevadensis F0078 ● ● Formica mlb04 micro grp F0085 ● ● Formica sp micro grp F0096 ● ● Formica sp micro grp F0087 ● ● Formica densiventris F0072 ● ● Formica talbotae F0156 monodomous ● ● Formica knighti F0015 ● ● Formica nepticula F0077 monodomous, polydomous ● ● Formica querquetulana F0095 ● ● monodomous, polydomous, supercolonial Formica querquetulana F0081

Supplementary Figure S3: Ancestral state reconstruction of nest structure using stochastic character mapping in Phy­ tools of Formica nest structure under all rates different model. Pie charts at nodes correspond to ancestral state estima­ tions and circles at tips correspond to states extant species. Numbers preceded by the letter F indicate extraction code and correspond to identifiers in Tables S1 and S2.

8 ● Polyergus rufescens F0149 ● ● Polyergus longicornis F0158 ● ● Polyergus vinosus F0124 ● ● Polyergus mexicanus F0153 ● Iberoformica subrufa D1135 ● ● Formica gagates F0120 ● Formica sp neog grp F0058 ● Formica obtusopilosa F0047 ● ● Formica lasioides F0040 ● ● ● ● Formica perpilosa F0039 ● ● Formica neogagates F0038 ● ● ● Formica manni F0037 ● Formica archboldi F0062 ● ● Formica pallidefulva F0066 ● ● Formica incerta F0059 ● ● Formica dolosa F0061 ● Formica moki F0129 ● ● Formica gnava F0105 ● ● Formica foreliana F0103 ● ● Formica gnava F0147 ● Formica subpolita F0137 ● Formica lemani F0130 ● ● ● Formica fuscocinerea F0115 ● ● Formica cinerea F0111 ● Formica cf xerophila F0140 ● ● ● Formica xerophila F0139 ● ● Formica subaenescens F0104 ● ● ● Formica accreta F0097 ● ● Formica transmontanis F0141 ● Formica aerata F0098 ● ● ● Formica argentea F0110 ● Formica nr occulta F0131 ● ● ● Formica montana nr F0117 ● Formica neoclara F0106 ● ● ● Formica cf altipetens F0100 ● ● ● Formica canadensis F0102 ● ● ● Formica altipetens F0099 ● Formica podzolica F0145 ● ● ● Formica glacialis F0121 ● Formica subsericea F0144 ● Formica cunicularia F0119 ● ● Formica rufibarbis F0136 ● ● Formica clara F0118 ● Formica candida F0155 ● ● Formica hewitti F0113 ● ● Formica neorufibarbis F0109 ● ● ● Formica neorufibarbis t F0126 ●● Formica neorufibarbis F0127 ● ● Formica neorufibarbis t F0108 ● ● Formica neorufibarbis F0128 ● Formica uralensis F0024 ● ● Formica exsecta F0034 ● ● ● Formica pressilabris F0033 ● ● Formica ulkei F0029 ● ● Formica exsectoides F0031 ● ● Formica exsectoides F0030 ● Formica sanguinea F0048 ● Formica aserva F0041 ● ● ● Formica rubicunda F0055 ● ● ● Formica pergandei F0053 ● ● Formica puberula F0122 ● ● ● Formica nr pergandei F0051 ● Formica emeryi F0050 ● ● Formica wheeleri F0057 ● ● ● Formica puberula F0056 ● ● Formica rubicunda F0054 ● Formica dakotensis F0069 ● ● Formica reflexa F0086 ● Formica truncorum F0023 ● ● Formica pratensis F0020 ● ● Formica lugubris F0021 ● ● Formica rufa F0018 ● ● ● Formica polyctena F0019 ● Formica obscuriventris F0011 ● ● Formica integra F0010 ● ● Formica cf fossaceps F0004 ● ● Formica obscuriventris F0006 ● ● ● Formica fossaceps F0003 ● ● Formica cf planipilis F0028 ● ● Formica ciliata F0001 ● ● Formica oreas F0013 ● ● Formica coloradensis F0002 ● ● Formica ravida F0026 ● ● Formica integroides F0114 ● ● Formica ravida F0027 ● ● ● Formica propinqua F0016 ● ● Formica integroides F0005 ● ● Formica cf subnitens F0014 ● ● ● Formica subnitens F0025 ● Formica obscuripes F0007 ● Formica sp micro grp F0157 ● Formica sp micro grp F0094 ● ● Formica sp micro grp F0092 ● ●● Formica sp micro grp F0074 ● ● Formica sp micro grp F0090 ●● Formica sp micro grp F0089 ●● Formica adamsi alpina F0068 ● ● Formica mlb01 micro grp F0082 ● ● Formica cf densiventris F0075 ● ● Formica nevadensis F0078 ● ● Formica mlb04 micro grp F0085 ● ● Formica sp micro grp F0096 ● ● Formica sp micro grp F0087 ● ● Formica densiventris F0072 ● ● Formica talbotae F0156 monogynous ● ● Formica knighti F0015 ● ● Formica nepticula F0077 monogynous, polygynous ● ● Formica querquetulana F0095 ● ● Formica querquetulana F0081

Supplementary Figure S4: Ancestral state reconstruction of colony structure using stochastic character mapping in Phytools of Formica colony structure under all rates different model. Pie charts at nodes correspond to ancestral state estimations and circles at tips correspond to states extant species. Numbers preceded by the letter F indicate extraction code and correspond to identifiers in Tables S1 and S2.

9 ● Polyergus rufescens F0149 ● ● Polyergus longicornis F0158 ● ● Polyergus vinosus F0124 ● ● Polyergus mexicanus F0153 ● Iberoformica subrufa D1135 ● Formica gagates F0120 ● ● Formica sp neog grp F0058 ● Formica obtusopilosa F0047 ● ● Formica lasioides F0040 ● ● ● Formica perpilosa F0039 ● ● Formica neogagates F0038 ● ● ● Formica manni F0037 ● ● Formica archboldi F0062 ● ● Formica pallidefulva F0066 ● ● Formica incerta F0059 ● ● Formica dolosa F0061 ● Formica moki F0129 ● ● Formica gnava F0105 ● ● Formica foreliana F0103 ● ● Formica gnava F0147 ● Formica subpolita F0137 ● Formica lemani F0130 ● ● ● Formica fuscocinerea F0115 ● ● Formica cinerea F0111 ● Formica cf xerophila F0140 ● ● ● Formica xerophila F0139 ● ● Formica subaenescens F0104 ● ● ● Formica accreta F0097 ● ● Formica transmontanis F0141 ● Formica aerata F0098 ● ● ● Formica argentea F0110 ● Formica nr occulta F0131 ● ● ● Formica montana nr F0117 ● Formica neoclara F0106 ● ● ● Formica cf altipetens F0100 ● ● ● Formica canadensis F0102 ● ● ● Formica altipetens F0099 ● ● Formica podzolica F0145 ● ● Formica glacialis F0121 ● Formica subsericea F0144 ● Formica cunicularia F0119 ● ● Formica rufibarbis F0136 ● ● Formica clara F0118 ● Formica candida F0155 ● ● Formica hewitti F0113 ● ● Formica neorufibarbis F0109 ● ● ● Formica neorufibarbis t F0126 ●● Formica neorufibarbis F0127 ● ● Formica neorufibarbis t F0108 ● ● Formica neorufibarbis F0128 ● Formica uralensis F0024 ● Formica exsecta F0034 ● ● Formica pressilabris F0033 ● Formica ulkei F0029 ● ● Formica exsectoides F0031 ● ● Formica exsectoides F0030 ● Formica sanguinea F0048 ● Formica aserva F0041 ● Formica rubicunda F0055 ● ● Formica pergandei F0053 ● ● Formica puberula F0122 ● ● ● Formica nr pergandei F0051 ● Formica emeryi F0050 ● ● Formica wheeleri F0057 ● ● ● Formica puberula F0056 ● Formica rubicunda F0054 ● Formica dakotensis F0069 ● Formica reflexa F0086 ● Formica truncorum F0023 ● Formica pratensis F0020 ● ● Formica lugubris F0021 ● ● Formica rufa F0018 ● ● Formica polyctena F0019 ● Formica obscuriventris F0011 ● ● Formica integra F0010 ● ● Formica cf fossaceps F0004 ● ● Formica obscuriventris F0006 ● Formica fossaceps F0003 ● Formica cf planipilis F0028 ● Formica ciliata F0001 ● Formica oreas F0013 ● Formica coloradensis F0002 ● Formica ravida F0026 ● Formica integroides F0114 ● Formica ravida F0027 ● Formica propinqua F0016 ● ● Formica integroides F0005 ● Formica cf subnitens F0014 ● Formica subnitens F0025 ● Formica obscuripes F0007 ● Formica sp micro grp F0157 ● Formica sp micro grp F0094 ● Formica sp micro grp F0092 ● ●● Formica sp micro grp F0074 ● ● Formica sp micro grp F0090 ● Formica sp micro grp F0089 ●● Formica adamsi alpina F0068 ● ● Formica mlb01 micro grp F0082 ● ● Formica cf densiventris F0075 ● ● Formica nevadensis F0078 ● ● Formica mlb04 micro grp F0085 ● ● Formica sp micro grp F0096 ● ● Formica sp micro grp F0087 ● ● Formica densiventris F0072 ● ● Formica talbotae F0156 independent colony founding ● ● Formica knighti F0015 ● Formica nepticula F0077 facultative temporary social parasitism, low freq. budding ● ● Formica querquetulana F0095 facultative temporary social parasitism, high freq. budding ● ● Formica querquetulana F0081 obligate temporary social parasitism, no budding (dulosis) permanent social parasitism

Supplementary Figure S5: Ancestral state reconstruction of colony founding mode using stochastic character mapping in Phytools of Formica colony founding under symmetrical rates model. Pie charts at the nodes correspond to ancestral state estimations and circles at tips correspond to states extant species. Numbers preceded by the letter F indicate extraction code and correspond to identifiers in Table S1 and S2.

10 Proformica_mongolica 1 Rossomyrmex_minuchae Polyergus_rufescens 1 Polyergus_longicornis 1 Polyergus_mexicanus 1 Polyergus_vinosus Iberoformica_subrufa 1 Formica_gagates 1 Formica_sp1_neog_grp Formica_obtusopilosa 1 Formica_lasioides 1 1 0.98 Formica_perpilosa 1 Formica_neogagates 1 1 Formica_manni Formica_archboldi 1 Formica_pallidefulva 1 Formica_incerta 1 1 Formica_dolosa Formica_moki 1 Formica_sp2_fusca_grp 1 Formica_gnava 1 Formica_foreliana 0.98 Formica_sp1_fusca_grp 1 Formica_gnava Formica_subpolita 1 1 Formica_sibylla 1 Formica_sibylla_cf Formica_cunicularia 1 Formica_orangea 0.87 Formica_rufibarbis 1 Formica_clara 1 Formica_occulta Formica_sp_fusca_grp 1 Formica_lemani 1 0.65 Formica_fuscocinerea 1 Formica_cinerea Formica_xerophila 0.96 1 0.97 Formica_xerophila_cf Formica_argentea 1 Formica_aerata 1 0.7 Formica_accreta 1 Formica_fusca_US 1 Formica_transmontanis 1 0.9 Formica_pacifica Formica_montana_nr 1 Formica_propatula 1 Formica_subsericea 0.92 1 Formica_subsericea_cf 1 1 Formica_glacialis Formica_neoclara 1 Formica_altipetens_cf 1 Formica_canadensis 1 Formica_altipetens Formica_candida 1 Formica_hewitti 1 Formica_neorufibarbis_t 1 Formica_neorufibarbis Formica_uralensis Formica_exsecta 1 1 Formica_pressilabris 0.53 0.74 1 Formica_mesasiatica Formica_ulkei 1 Formica_opaciventris 1 Formica_exsectoides Formica_reflexa 1 Formica_dakotensis 1 0.36 Formica_sanguinea Formica_aserva 1 Formica_rubicunda2 1 1 Formica_pergandei3 1 Formica_pergandei2 1 1 Formica_puberula2 Formica_emeryi Formica_pergandei1 1 1 Formica_rubicunda1 1 1 Formica_puberula1 1 Formica_wheeleri Formica_talbotae Formica_truncorum 1 Formica_frontalis_cf 1 Formica_lugubris 0.49 Formica_pratensis 0.71 Formica_polyctena 1 1 Formica_rufa Formica_planipilis 1 Formica_oreas 1 0.7 Formica_ciliata Formica_obscuriventris2 1 1 Formica_fossaceps 0.82 Formica_obscuriventris1 0.99 Formica_interga 1 1 Formica_fossaceps_cf Formica_subnitens 1 Formica_obscuripes 0.87 Formica_coloradensis 1 Formica_ravida2 1 0.73 Formica_integroides2 1 Formica_ravida1 1 Formica_integroides1 1 Formica_propinqua Formica_sp12_micro_grp Formica_adamsi_alpina 1 1 Formica_sp11_micro_grp Formica_sp8_micro_grp 1 1 Formica_sp7_micro_grp 0.96 Formica_nevadensis 0.75 1 Formica_sp1_micro_grp Formica_sp3_micro_grp Formica_sp4_micro_grp 0.78 1 Formica_sp5_micro_grp 0.49 Formica_knighti 1 Formica_nepticula 1 Formica_querquetulana

0.8

Supplementary Figure S6: Species tree inferred under coalescent using ASTRAL­III and 1,733 weighted supergene trees identified using statistical binning pipeline. Support is expressed in local posterior probability and branch lengths are in coalescent units.

11 Proformica_mongolica 1 Rossomyrmex_minuchae Polyergus_rufescens 1 Polyergus_longicornis 0.99 Polyergus_mexicanus 1 Polyergus_vinosus Iberoformica_subrufa 1 Formica_gagates 0.99 Formica_sp1_neog_grp Formica_obtusopilosa 1 Formica_lasioides 1 1 1 Formica_perpilosa 1 Formica_neogagates 1 1 Formica_manni Formica_archboldi 1 Formica_pallidefulva 1 Formica_incerta 1 1 Formica_dolosa Formica_moki 1 Formica_sp2_fusca_grp 1 Formica_gnava 1 Formica_foreliana 0.81 Formica_sp1_fusca_grp 1 Formica_gnava Formica_subpolita 1 1 Formica_sibylla 1 Formica_sibylla_cf Formica_cunicularia 1 Formica_orangea 0.97 Formica_rufibarbis 1 Formica_clara 1 Formica_occulta Formica_fuscocinerea 1 Formica_cinerea 1 1 Formica_lemani 0.63 Formica_sp_fusca_grp Formica_xerophila 0.96 1 0.99 Formica_xerophila_cf Formica_argentea 1 Formica_aerata 10.72 Formica_accreta 1 Formica_fusca_US 1 Formica_transmontanis 1 0.69 Formica_pacifica Formica_montana_nr 1 Formica_propatula 1 Formica_subsericea 0.49 0.98 Formica_subsericea_cf 1 0.96 Formica_glacialis Formica_neoclara 1 Formica_altipetens_cf 1 Formica_canadensis 1 Formica_altipetens Formica_candida 1 Formica_hewitti 1 Formica_neorufibarbis_t 1 Formica_neorufibarbis Formica_uralensis Formica_exsecta 1 1 Formica_pressilabris 0.76 0.99 1 Formica_mesasiatica Formica_ulkei 1 Formica_opaciventris 1 Formica_exsectoides Formica_sanguinea Formica_aserva 1 1 Formica_rubicunda2 1 0.99 Formica_pergandei3 1 Formica_pergandei2 1 1 Formica_puberula2 Formica_emeryi Formica_puberula1 1 0.93 Formica_wheeleri 0.99 1 Formica_pergandei1 1 Formica_rubicunda1 Formica_reflexa 1 Formica_dakotensis Formica_talbotae Formica_lugubris 0.6 Formica_truncorum 1 1 Formica_frontalis_cf 0.97 Formica_pratensis 0.97 Formica_polyctena 1 1 Formica_rufa Formica_subnitens 1 Formica_obscuripes 0.85 Formica_coloradensis 1 Formica_ravida2 1 0.96 Formica_integroides2 0.92 Formica_ravida1 1 Formica_integroides1 1 1 Formica_propinqua Formica_planipilis 1 Formica_oreas 1 0.49 Formica_ciliata Formica_obscuriventris2 1 1 Formica_fossaceps 0.69 Formica_obscuriventris1 0.8 Formica_fossaceps_cf 1 Formica_interga Formica_sp12_micro_grp Formica_adamsi_alpina 1 1 Formica_sp11_micro_grp Formica_nevadensis 0.54 1 Formica_sp1_micro_grp 0.93 Formica_sp8_micro_grp 0.99 Formica_sp7_micro_grp 0.45 Formica_sp4_micro_grp 1 1 Formica_sp5_micro_grp Formica_sp3_micro_grp 0.59 Formica_knighti 1 Formica_nepticula 1 Formica_querquetulana

0.8

Supplementary Figure S7: Species tree inferred under coalescent using ASTRAL­III and all 2,242 individual UCE locus trees. Support is expressed in local posterior probability and branch lengths are in coalescent units.

12 Proformica_mongolica 1 Rossomyrmex_minuchae Polyergus_longicornis 1 Polyergus_rufescens 0.59 Polyergus_mexicanus 0.91 Polyergus_vinosus Iberoformica_subrufa 1 Formica_sp1_neog_grp 0.54 Formica_obtusopilosa Formica_lasioides 0.98 0.31 Formica_perpilosa 1 Formica_neogagates 0.97 1 0.98 Formica_manni Formica_dolosa 1 Formica_incerta 0.52 Formica_archboldi 0.99 Formica_pallidefulva 1 Formica_gagates Formica_moki Formica_subpolita 0.48 0.93 Formica_sibylla 1 0.47 Formica_sibylla_cf Formica_gnava 1 Formica_gnava 0.85 Formica_sp2_fusca_grp 0.4 0.68 Formica_sp1_fusca_grp 0.37 Formica_foreliana Formica_orangea 1 Formica_cunicularia 0.7 Formica_rufibarbis 0.96 Formica_clara Formica_fuscocinerea 0.44 0.77 Formica_cinerea 0.81 0.99 Formica_lemani 0.45 Formica_sp_fusca_grp Formica_xerophila 0.85 1 Formica_xerophila_cf Formica_occulta 0.48 Formica_transmontanis 1 Formica_pacifica 0.36 0.44 Formica_argentea 0.69 Formica_aerata Formica_accreta 0.77 0.94 Formica_fusca_US 0.27 Formica_montana_nr 0.94 0.47 Formica_propatula 0.49 Formica_subsericea Formica_subsericea_cf 0.7 1 Formica_glacialis 0.59 Formica_neoclara 0.53 Formica_altipetens_cf 0.56 Formica_canadensis 0.91 Formica_altipetens Formica_candida 1 Formica_neorufibarbis 0.46 Formica_hewitti 0.44 Formica_neorufibarbis_t Formica_ulkei 1 Formica_opaciventris 0.83 0.51 Formica_exsectoides 0.45 Formica_uralensis 0.37 Formica_pressilabris 1 Formica_exsecta 0.66 Formica_mesasiatica Formica_aserva 0.45 Formica_sanguinea 0.86 0.84 Formica_rubicunda2 Formica_pergandei3 0.54 0.96 Formica_pergandei2 1 0.4 Formica_puberula2 Formica_emeryi 0.54 Formica_puberula1 0.72 Formica_wheeleri 0.43 Formica_pergandei1 0.58 0.61 Formica_rubicunda1 Formica_reflexa 1 Formica_dakotensis Formica_lugubris Formica_truncorum 1 1 Formica_frontalis_cf 0.43 Formica_pratensis 0.37 0.73 Formica_polyctena 0.96 Formica_rufa Formica_subnitens 0.66 Formica_obscuripes Formica_obscuriventris2 0.98 Formica_interga 0.68 0.97 0.83 Formica_fossaceps_cf 0.42 Formica_fossaceps 0.47 0.35 Formica_obscuriventris1 Formica_ciliata 0.98 Formica_oreas 0.5 Formica_planipilis 0.42 0.58 Formica_coloradensis Formica_ravida2 0.660.44 Formica_integroides2 0.37 Formica_ravida1 0.99 Formica_integroides1 0.53 Formica_propinqua Formica_talbotae 0.29 Formica_sp12_micro_grp Formica_sp3_micro_grp 10.35 Formica_sp8_micro_grp 0.59 Formica_sp7_micro_grp 0.34 Formica_knighti Formica_nepticula 0.63 0.39Formica_querquetulana 0.42 Formica_sp11_micro_grp 0.57 Formica_sp4_micro_grp 0.46 0.53 Formica_sp5_micro_grp Formica_adamsi_alpina 0.39 Formica_nevadensis 0.42 Formica_sp1_micro_grp

0.6

Supplementary Figure S8: Species tree inferred under coalescent using ASTRAL­III and 67 individual UCE locus trees for which at least 50 % of sequence length was available for Formica talbotae. Support is expressed in local posterior probability and branch lengths are in coalescent units.

13 Proformica_mongolica_F0150 1/NA/1 Rossomyrmex_minuchae_F0152 Polyergus_rufescens_F0149 1/NA/1 Polyergus_longicornis_F0158 0.96/0.67/1 Polyergus_mexicanus_F0154 0.97/0/1 Polyergus_mexicanus_F0153 0.93/1/0.99 Polyergus_vinosus_F0124 Iberoformica_subrufa_D1135 Formica_gagates_F0120 1/NA/1 Formica_sp_neog_grp_F0058 0.91/0.67/0.99 Formica_obtusopilosa_F0046 1/NA/1 1/NA/1 Formica_obtusopilosa_F0047 Formica_lasioides_F0036 1/NA/1 Formica_lasioides_F0040 1/NA/1 0.81/0.76/0.99 1/NA/1 Formica_perpilosa_F0039 1/NA/1 Formica_manni_F0037 1/NA/1 Formica_neogagates_F0038 1/NA/1 Formica_archboldi_F0062 0.97/1/1 Formica_pallidefulva_F0060 1/NA/1 Formica_pallidefulva_F0066 1/NA/1 Formica_dolosa_F0061 0.92/0/1 Formica_dolosa_F0063 0.91/0.67/1 1/NA/1 Formica_incerta_F0059 0.93/0.5/1 Formica_incerta_F0065 Formica_moki_F0129 1/NA/1 Formica_sp_fusca_grp_F0161 1/NA/1 Formica_gnava_F0105 0.83/0.71/1 Formica_foreliana_F0103 0.5/0.17/0.97 Formica_ _F0147 0.72/0.15/0.99 gnava Formica_sp_fusca_grp_F0160 Formica_sibylla_F0132 1/NA/1 Formica_cf_sibylla_F0133 1/NA/1 Formica_subpolita_F0137 1/NA/1 Formica_subpolita_F0138 1/NA/1 Formica_sp_fusca_grp_F0101 1/NA/1 Formica_lemani_F0130 0.94/0/1 Formica_cinerea_F0111 1/NA/1 Formica_fuscocinerea_F0115 Formica_xerophila_F0139 1/NA/1 1/NA/1 Formica_cf_xerophila_F0140 Formica_pacifica_F0135 1/NA/1 Formica_transmontanis_F0141 0.98/0/1 1/NA/1 Formica_accreta_F0097 0.73/0.21/0.99 1/NA/1 Formica_subaenescens_F0104 Formica_aerata_F0098 -0.33/0.14/0.98 1/NA/1 Formica_sp_fusca_grp_F0165 1/NA/1 Formica_argentea_F0110 0.94/0/0.99 Formica_sp_fusca_grp_F0143 -0.57/0.005/0.98 Formica_nr_occulta_F0131 1/NA/1 Formica_montana_nr_F0117 1/NA/1 Formica_propatula_F0116 Formica_neoclara_F0106 0.94/0.8/1 0.96/0.67/1 Formica_cf_altipetens_F0100 1/NA/1 Formica_altipetens_F0099 0.58/0.051/0.96 0.9/0.67/1 0.7/0.042/0.99 Formica_canadensis_F0102 Formica_glacialis_F0121 0.96/0.67/1 Formica_podzolica_F0145 1/NA/1 Formica_subsericea_F0146 1/NA/1 Formica_subsericea_F0159 0.41/0.5/0.94 Formica_subsericea_F0144 Formica_orangea_F0134 1/NA/1 Formica_cunicularia_F0119 1/NA/1 Formica_clara_F0118 1/NA/1 Formica_rufibarbis_F0136 Formica_candida_F0155 Formica_hewitti_F0113 0.98/0/1 1/NA/1 Formica_sp_fusca_grp_F0142 Formica_neorufibarbis_F0109 0.11/0/0.97 0.96/0.67/1 -0.55/0.029/0.99 Formica_neorufibarbis_F0107 1/NA/1 Formica_neorufibarbis_F0128 0.57/0.038/0.99 Formica_neorufibarbis_t_F0108 0.98/0/1 Formica_neorufibarbis_F0127 0.94/0/1 Formica_neorufibarbis_t_F0126 Formica_uralensis_F0024 Formica_exsecta_F0034 0.98/0/1 1/NA/1 Formica_mesasiatica_F0035 0.41/0.054/0.97 0.57/0.083/0.97 0.98/0/1 Formica_pressilabris_F0033 Formica_ulkei_F0029 1/NA/1 Formica_exsectoides_F0031 0.97/0/1 Formica_exsectoides_F0032 0.5/0/0.99 Formica_exsectoides_F0030 Formica_sanguinea_F0048 1/NA/1 Formica_sanguinea_F0049 Formica_aserva_F0043 Formica_aserva_F0041 1/NA/1 0.48/0.018/0.99 1/NA/1 1/NA/1 Formica_aserva_F0042 1/NA/1 Formica_aserva_F0163 -0.15/0.31/0.97 Formica_aserva_F0044 0.93/1/0.99 0.89/0.55/1 Formica_aserva_F0067 Formica_rubicunda_F0055 0.54/0/0.99 Formica_pergandei_F0053 1/NA/1 Formica_nr_pergandei_F0051 1/NA/1 1/NA/1 Formica_puberula_F0122 Formica_emeryi_F0050 Formica_sp_sang_grp_F0123 1/NA/1 1/NA/1 Formica_rubicunda_F0054 1/NA/1 0.85/1/0.98 Formica_puberula_F0056 1/NA/1 Formica_wheeleri_F0057 Formica_dakotensis_F0071 1/NA/1 Formica_dakotensis_F0069 0.93/0.5/1 Formica_dakotensis_F0070 0.86/0/1 Formica_reflexa_F0086 Formica_cf_frontalis_F0017 0.93/1/1 Formica_truncorum_F0023 1/NA/1 Formica_pratensis_F0020 0.92/0.86/1 Formica_lugubris_F0021 0.35/0.015/0.98 Formica_polyctena_F0019 1/NA/1 0.44/0.47/0.96 Formica_rufa_F0018 Formica_cf_planipilis_F0028 1/NA/1 Formica_ciliata_F0001 0.94/0.8/1 Formica_oreas_F0013 0.92/0/1 Formica_oreas_F0012 0.87/1/1 0.048/0.7/0.98 Formica_oreas_F0125 Formica_obscuriventris_F0011 Formica_fossaceps_F0003 0.95/0/1 0.024/0/0.99 1/NA/1 Formica_obscuriventris_F0006 0.041/0.67/0.98 Formica_cf_fossaceps_F0004 0.97/0/1 1/NA/1 Formica_cf_interga_F0010 Formica_coloradensis_F0002 1/NA/1 Formica_ravida_F0026 0.38/0.11/0.98 Formica_integroides_F0114 0.96/0.67/1 Formica_ravida_F0027 0.97/0/1 Formica_integroides_F0005 1/NA/1 0.35/0.77/1 Formica_propinqua_F0016 Formica_cf_subnitens_F0014 1/NA/1 Formica_subnitens_F0022 1/NA/1 0.83/0/1 0.95/0/1 Formica_subnitens_F0025 Formica_obscuripes_F0007 1/NA/1 Formica_obscuripes_F0008 0.97/0/1 Formica_obscuripes_F0009 0.94/0/1 Formica_obscuripes_F0162 Formica_sp_micro_grp_F0157 Formica_sp_micro_grp_F0094 Formica_sp_micro_grp_F0074 0.96/0/1 0.94/0.4/1 Formica_sp_micro_grp_F0092 0.92/0/1 1/NA/1 Formica_sp_micro_grp_F0090 0.94/0.4/1 Formica_adamsi_alpina_F0068 -0.73/0/1 Formica_sp_micro_grp_F0089 Formica_sp_micro_grp_F0087 0.95/0.5/1 0.78/0/1 Formica_sp_micro_grp_F0096 0.45/0.53/0.99 Formica_densiventris_F0073 0.4/0.051/0.99 Formica_densiventris_F0072 0.94/0/1 0.24/0.31/0.98 Formica_cf_densiventris_F0164 Formica_talbotae_F0156 -0.44/0.052/0.97 Formica_knighti_F0015 0.46/0.67/0.95 Formica_nepticula_F0077 0.55/0.88/0.98 1/NA/1 Formica_querquetulana_F0081 0.9/0.67/1 Formica_querquetulana_F0095 Formica_cf_densiventris_F0075 0.89/0.91/1 Formica_mlb01_micro_grp_F0082 Formica_mlb02_micro_grp_F0083 0.072/0.87/0.98 0.94/0.8/1 Formica_mlb03_micro_grp_F0084 0.9/0.33/1 Formica_mlb04_micro_grp_F0085 0.28/0.17/0.94 0.74/0.94/1 Formica_sp_micro_grp_F0091 Formica_nevadensis_F0080 0.35/0.83/0.99 Formica_nevadensis_F0078 0.94/0/0.99 Formica_nevadensis_F0079 0.9/0.67/1 Formica_sp_micro_grp_F0093

Supplementary Figure S9: Support from quartet sampling analyses. The values are: quartet concordance score (QC) / quartet differential score (QD) / quartet informativeness score (QI). Briefly, QC measures how often the concordant quartet topologies are inferred over discordant quartets in different replicates, with a value equal 1 when all replicates result in concordant topology, QD measures whether frequencies of two discordant topologies are equal (=1) or skewed (<1), and QI measures what proportion of replicates were informative. Contrast values for colony dependent clade or monophyly of difficilis group (1/NA/1), meaning that all replicates were informative and resulted in the same topology with support for placement of clade of Palearctic species related to Formica rufibarbis (­0.55/0.029/0.99) indicating that discordant topologies were inferred more often than those concordant with the input tree, that quartet topologies were highly skewed, but most replicates were informative. The former indicates maximal support, while the latter shows that the position of this clade within the ”fusca grade” is highly uncertain, perhaps due to incomplete lineage sorting.

14 Supplementary Table S1: Voucher specimens used in this study. Detailed collection data can be accessed at Zenodo (DOI: 10.5281/zenodo.4341310).

Extraction ID Taxon Specimen ID Country Latitude Longitude D1135 Iberoformica subrufa ASU­SIBR2148 ES 37.185 ­3.485 F0001 Formica ciliata ASU­SIBR2001 US 38.26117 ­112.51433 F0002 Formica coloradensis ASU­SIBR2002 US 38.42233 ­109.18017 F0003 Formica fossaceps ASU­SIBR2003 US 38.39467 ­109.165 F0004 Formica cf fossaceps ASU­SIBR2004 CA 44.53050 ­64.31912 F0005 Formica integroides ASU­SIBR2005 US 38.67525 ­119.99428 F0006 Formica obscuriventris ASU­SIBR2006 US 39.12017 ­107.44767 F0007 Formica obscuripes ASU­SIBR2007 US 48.1728 ­122.6714 F0008 Formica obscuripes ASU­SIBR2008 US 38.23367 ­112.44917 F0009 Formica obscuripes ASU­SIBR2009 US 42.44788 ­84.01688 F0010 Formica cf interga ASU­SIBR2010 CA 44.68839 ­63.66808 F0011 Formica obscuriventris ASU­SIBR2011 US 42.45103 ­84.01878 F0012 Formica oreas ASU­SIBR2012 US 38.67577 ­119.99431 F0013 Formica oreas ASU­SIBR2013 US 38.84233 ­106.53817 F0014 Formica cf subnitens ASU­SIBR2014 US 38.67757 ­119.98343 F0015 Formica knighti ASU­SIBR2015 US 41.8652 ­70.64777 F0016 Formica propinqua ASU­SIBR2016 US 39.44008 ­120.32349 F0017 Formica cf frontalis ASU­SIBR2017 KG 42.8 77.47 F0018 Formica rufa ASU­SIBR2018 PL 51.155 16.985 F0019 Formica polyctena ASU­SIBR2019 PL 51.39 17.5 F0020 Formica pratensis ASU­SIBR2020 PL 51.3867 16.805 F0021 Formica lugubris ASU­SIBR2021 IT 46.586 12.105 F0022 Formica subnitens ASU­SIBR2022 US 39.82308 ­120.13748 F0023 Formica truncorum ASU­SIBR2023 PL 50.33 16.55 F0024 Formica uralensis ASU­SIBR2024 PL 51.38 23.55 F0025 Formica subnitens ASU­SIBR2025 US 39.591 ­108.818 F0026 Formica ravida ASU­SIBR2026 US 39.1695 ­107.94817 F0027 Formica ravida ASU­SIBR2027 US 39.43221 ­120.24265 F0028 Formica cf planipilis ASU­SIBR2028 US 40.72538 ­120.20708 F0029 Formica ulkei ASU­SIBR2029 CA 44.49808 ­63.92692 F0030 Formica exsectoides ASU­SIBR2030 US 41.87383 ­70.65183 F0031 Formica exsectoides ASU­SIBR2031 US 38.73367 ­104.896 F0032 Formica exsectoides ASU­SIBR2032 US 43.01548 ­77.57365 F0033 Formica pressilabris ASU­SIBR2033 SE 68.3381 18.7639 F0034 Formica exsecta ASU­SIBR2034 PL 50.577 22.984 F0035 Formica mesasiatica ASU­SIBR2035 KG 42.33 78.24 F0036 Formica lasioides ASU­SIBR2036 US 33.18522 ­116.28849 F0037 Formica manni ASU­SIBR2037 US 41.63632 ­119.84183 F0038 Formica neogagates ASU­SIBR2038 US 39.42998 ­120.24087 F0039 Formica perpilosa ASU­SIBR2039 US 31.83945 ­109.03615 F0040 Formica lasioides ASU­SIBR2040 US 39.43163 ­120.24059 F0041 Formica aserva ASU­SIBR2041 US 37.68319 ­119.17078 F0042 Formica aserva ASU­SIBR2042 US 48.66648 ­122.96887 F0043 Formica aserva ASU­SIBR2043 CA 44.46380 ­63.58094 F0044 Formica aserva ASU­SIBR2044 US 38.81400 ­106.28217 F0046 Formica obtusopilosa ASU­SIBR2046 US 38.71217 ­111.94833 F0047 Formica obtusopilosa ASU­SIBR2047 US 39.50633 ­108.76217 F0048 Formica sanguinea ASU­SIBR2048 PL 50.448 19.995 F0049 Formica sanguinea ASU­SIBR2049 KG 42.67 77.18 F0050 Formica emeryi ASU­SIBR2050 US 39.03167 ­104.79633 F0051 Formica nr pergandei ASU­SIBR2051 US 39.21529 ­121.04322 F0053 Formica pergandei ASU­SIBR2053 US 42.45278 ­84.01939 F0054 Formica rubicunda ASU­SIBR2054 US 39.03033 ­105.44217 F0055 Formica rubicunda ASU­SIBR2055 US 42.45919 ­84.01429 F0056 Formica puberula ASU­SIBR2056 US 37.88800 ­109.45433 F0057 Formica wheeleri ASU­SIBR2057 US 31.91433 ­109.271 F0058 Formica sp neog grp ASU­SIBR2058 US 37.84367 ­109.36967 F0059 Formica incerta ASU­SIBR2059 US 42.45151 ­84.01901 F0060 Formica pallidefulva ASU­SIBR2060 US 42.45948 ­84.0256 F0061 Formica dolosa ASU­SIBR2061 US 30.35995 ­84.41848 F0062 Formica archboldi ASU­SIBR2062 US 30.35995 ­84.41848 F0063 Formica dolosa ASU­SIBR2063 US 41.81444 ­70.66315 F0065 Formica incerta ASU­SIBR2065 US 41.8652 ­70.64777

15 Supplementary Table S1: Voucher specimens used in this study, continued.

Extraction ID Taxon Specimen ID Country Latitude Longitude F0066 Formica pallidefulva ASU­SIBR2066 US 30.2275 ­98.1788 F0067 Formica aserva ASU­SIBR2067 US 38.66283 ­111.94117 F0068 Formica adamsi alpina ASU­SIBR2068 US 38.39317 ­107.19683 F0069 Formica dakotensis ASU­SIBR2069 US 34.02867 ­109.1855 F0070 Formica dakotensis ASU­SIBR2070 US 38.63500 ­111.94817 F0071 Formica dakotensis ASU­SIBR2071 US 39.41344 ­120.32162 F0072 Formica densiventris ASU­SIBR2072 US 32.65729 ­109.85841 F0073 Formica densiventris ASU­SIBR2073 US 39.75867 ­108.78783 F0074 Formica sp micro grp ASU­SIBR2074 US 38.64883 ­111.94983 F0075 Formica cf densiventris ASU­SIBR2075 US 38.21541 ­119.74609 F0077 Formica nepticula ASU­SIBR2077 US 41.87367 ­70.65233 F0078 Formica nevadensis ASU­SIBR2078 US 39.35788 ­122.74716 F0079 Formica nevadensis ASU­SIBR2079 US 41.56204 ­123.21177 F0080 Formica nevadensis ASU­SIBR2080 US 39.41318 ­120.32038 F0081 Formica querquetulana ASU­SIBR2081 US 41.87383 ­70.65183 F0082 Formica mlb01 micro grp ASU­SIBR2082 US 37.21317 ­118.64671 F0083 Formica mlb02 micro grp ASU­SIBR2083 US 40.94628 ­123.0427 F0084 Formica mlb03 micro grp ASU­SIBR2084 US 39.29111 ­120.67918 F0085 Formica mlb04 micro grp ASU­SIBR2085 US 39.40022 ­120.55818 F0086 Formica reflexa ASU­SIBR2086 CA 44.53050 ­64.31912 F0087 Formica sp micro grp ASU­SIBR2087 US 38.42033 ­107.62783 F0089 Formica sp micro grp ASU­SIBR2089 US 38.18833 ­107.62067 F0090 Formica sp micro grp ASU­SIBR2090 US 38.83800 ­106.56167 F0091 Formica sp micro grp ASU­SIBR2091 US 38.83800 ­106.56167 F0092 Formica sp micro grp ASU­SIBR2092 US 38.64983 ­111.95083 F0093 Formica sp micro grp ASU­SIBR2093 US 41.56204 ­123.21177 F0094 Formica sp micro grp ASU­SIBR2094 US 38.838 ­106.56167 F0095 Formica querquetulana ASU­SIBR2095 US 41.87167 ­70.651 F0096 Formica sp micro grp ASU­SIBR2096 US 39.04167 ­104.6615 F0097 Formica accreta ASU­SIBR2097 US 39.30805 ­120.66831 F0098 Formica aerata ASU­SIBR2098 US 37.80145 ­118.5299 F0099 Formica altipetens ASU­SIBR2099 US 33.90683 ­109.1245 F0100 Formica cf altipetens ASU­SIBR2100 US 38.6739 ­119.99411 F0101 Formica sp fusca grp ASU­SIBR2101 US 37.38932 ­118.76612 F0102 Formica canadensis ASU­SIBR2102 US 39.12017 ­107.44 F0103 Formica foreliana ASU­SIBR2103 US 31.43100 ­111.16967 F0104 Formica subaenescens ASU­SIBR2104 US 37.92051 ­122.57471 F0105 Formica gnava ASU­SIBR2105 US 32.64983 ­109.81717 F0106 Formica neoclara ASU­SIBR2106 US 39.50332 ­120.24157 F0107 Formica neorufibarbis ASU­SIBR2107 US 38.71467 ­106.223 F0108 Formica neorufibarbis t ASU­SIBR2108 US 36.67239 ­118.34463 F0109 Formica neorufibarbis ASU­SIBR2109 US 39.42505 ­120.24285 F0110 Formica argentea ASU­SIBR2110 MX 31.00261 ­115.5497 F0111 Formica cinerea ASU­SIBR2111 PL 51.39 17.5 F0113 Formica hewitti ASU­SIBR2113 US 38.67984 ­119.98373 F0114 Formica integroides ASU­SIBR2114 US 40.11490 ­120.32086 F0115 Formica fuscocinerea ASU­SIBR2115 DE 48.16452 11.50075 F0116 Formica propatula ASU­SIBR2116 MX 17.1873 ­96.62085 F0117 Formica montana nr ASU­SIBR2117 MX 17.61752 ­96.36799 F0118 Formica clara ASU­SIBR2118 KG 42.66 77.47 F0119 Formica cunicularia ASU­SIBR2119 PL 51.175 17.068 F0120 Formica gagates ASU­SIBR2120 CZ 48.869 16.652 F0121 Formica glacialis ASU­SIBR2121 CA 44.98735 ­64.06002 F0122 Formica puberula ASU­SIBR2122 US 45.0446 ­110.68078 F0123 Formica sp sang grp ASU­SIBR2123 US 45.13217 ­111.06283 F0124 Polyergus vinosus ASU­SIBR2124 US 32.85323 ­116.43738 F0125 Formica oreas ASU­SIBR2125 US 38.05273 ­119.32138 F0126 Formica neorufibarbis t ASU­SIBR2126 US 38.06165 ­119.34966 F0127 Formica neorufibarbis ASU­SIBR2127 US 38.06207 ­119.34888 F0128 Formica neorufibarbis ASU­SIBR2128 US 38.04167 ­119.29781 F0129 Formica moki ASU­SIBR2129 US 38.9 ­121.02 F0130 Formica lemani ASU­SIBR2130 PL 50.33 16.55

16 Supplementary Table S1: Voucher specimens used in this study, continued.

Extraction ID Taxon Specimen ID Country Latitude Longitude F0131 Formica nr occulta ASU­SIBR2131 US 31.91375 ­109.26757 F0132 Formica sibylla ASU­SIBR2132 US 39.43221 ­120.24258 F0133 Formica cf sibylla ASU­SIBR2133 US 39.431 ­120.331 F0134 Formica orangea ASU­SIBR2134 KG 42.6 75.85 F0135 Formica pacifica ASU­SIBR2135 US 48.1728 ­122.6714 F0136 Formica rufibarbis ASU­SIBR2136 PL 52.76917 14.30778 F0137 Formica subpolita ASU­SIBR2137 US 39.50444 ­120.23817 F0138 Formica subpolita ASU­SIBR2138 US 38.72450 ­112.20717 F0139 Formica xerophila ASU­SIBR2139 US 35.80350 ­112.11767 F0140 Formica cf xerophila ASU­SIBR2140 US 38.85876 ­122.41834 F0141 Formica transmontanis ASU­SIBR2141 US 39.188 ­123.757 F0142 Formica sp fusca grp ASU­SIBR2142 US 38.83767 ­106.55933 F0143 Formica sp fusca grp ASU­SIBR2143 US 39.21533 ­121.04019 F0144 Formica subsericea ASU­SIBR2144 US 35.695 ­78.69688 F0145 Formica podzolica ASU­SIBR2145 US 43.01616 ­77.57632 F0146 Formica subsericea ASU­SIBR2146 US 42.45103 ­84.01885 F0147 Formica gnava ASU­SIBR2147 US 31.93200 ­109.17783 F0149 Polyergus rufescens ASU­SIBR2150 KG 42.67 77.18 F0150 Proformica mongolica ASU­SIBR2151 KG 42.77 74.66 F0152 ASU­SIBR2153 ES 36.89 ­2.78 F0153 Polyergus mexicanus ASU­SIBR2154 US 39.41591 ­120.31704 F0154 Polyergus mexicanus ASU­SIBR2155 US 31.90882 ­109.25211 F0155 Formica candida ASU­SIBR2156 KG 42.08 76.73 F0156 Formica talbotae ASU­SIBR2157 US 43.38 ­95.185 F0157 Formica sp micro grp ASU­SIBR2158 US 42.45947 ­84.02559 F0158 Polyergus longicornis ASU­SIBR2159 US 30.35995 ­84.41848 F0159 Formica subsericea ASU­SIBR2160 US 38.9879 ­77.2495 F0160 Formica sp fusca grp ASU­SIBR2161 US 31.8729 ­109.23504 F0161 Formica sp fusca grp ASU­SIBR2162 US 32.59105 ­107.97398 F0162 Formica obscuripes ASU­SIBR2163 US 48.03381 ­110.22798 F0163 Formica aserva ASU­SIBR2164 US 46.7059 ­114.53617 F0164 Formica cf densiventris ASU­SIBR2165 US 38.10233 ­111.33653 F0165 Formica sp fusca grp ASU­SIBR2166 US 48.52302 ­113.3808

17 Supplementary Table S2: Life history of Formica and other taxa used in this study. Abbreviations: ASR: ancestral state reconstruction; ICF: independent colony founding; DCF: dependent colony founding; TSP: temporary social parasitism.

Extraction ID Taxon Species group Life history Colony structure Nest structure Colony founding ASR colony founding References F0097 Formica accreta “fusca” free-living monogynous, polygynous monodomous ICF independent colony founding Francoeur 1973; Frumhoff and Ward 1992; JT Longino and PS Ward unpubl. collection data F0068 Formica adamsi alpina difficilis free-living, TSP monogynous, polygynous monodomous TSP, DCF: budding facultative temporary parasitism, low frequency budding SP Cover unpubl. collection data F0098 Formica aerata “fusca” free-living monogynous, polygynous monodomous, polydomous ICF independent colony founding Francoeur 1973; PS Ward unpubl. collection data F0099 Formica altipetens “fusca” free-living monogynous, polygynous monodomous, polydomous ICF, DCF: budding independent colony founding SP Cover unpubl. collection data F0100 Formica cf altipetens “fusca” free-living monogynous, polygynous monodomous, polydomous ICF, DCF: budding independent colony founding SP Cover unpubl. collection data F0062 Formica archboldi pallidevulva free-living monogynous, polygynous monodomous ICF: haplometrosis, pleometrosis independent colony founding Trager et al. 2007, SP Cover unpubl. collection data F0110 Formica argentea “fusca” free-living monogynous, polygynous monodomous ICF: haplometrosis, pleometrosis independent colony founding Snyder 1992; SP Cover unpubl. collection data F0041 Formica aserva sanguinea free-living, TSP, fac. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Savolainen and Deslippe 2001; SP Cover unpubl. collection data F0042 Formica aserva sanguinea free-living, TSP, fac. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Savolainen and Deslippe 2001; SP Cover unpubl. collection data F0043 Formica aserva sanguinea free-living, TSP, fac. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Savolainen and Deslippe 2001; SP Cover unpubl. collection data F0044 Formica aserva sanguinea free-living, TSP, fac. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Savolainen and Deslippe 2001; SP Cover unpubl. collection data F0067 Formica aserva sanguinea free-living, TSP, fac. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Savolainen and Deslippe 2001; SP Cover unpubl. collection data F0163 Formica aserva sanguinea free-living, TSP, fac. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Savolainen and Deslippe 2001; SP Cover unpubl. collection data F0102 Formica canadensis “fusca” free-living monogynous, polygynous monodomous, polydomous ICF, DFC: budding independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0155 Formica candida “fusca” free-living monogynous, polygynous monodomous, polydomous ICF independent colony founding Seifert 2004, Seifert 2018 F0001 Formica ciliata integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP, DCF: budding facultative temporary parasitism, high frequency budding Wheeler 1913; SP Cover unpubl. collection data F0111 Formica cinerea “fusca” free-living monogynous, polygynous monodomous, polydomous, supercolonial ICF: haplometrosis, pleometrosis, DCF: budding independent colony founding Zhu et al. 2003; Lindström et al. 1996; Goropashnaya et al. 2001 F0118 Formica clara “fusca” free-living monogynous, polygynous monodomous ICF independent colony founding Seifert 2007; Seifert 2018; Seifert and Schultz 2009 F0002 Formica coloradensis integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP, DCF: budding facultative temporary parasitism, high frequency budding Creighton 1940; SP Cover unpubl. collection data F0119 Formica cunicularia “fusca” free-living monogynous, polygynous monodomous ICF: haplometrosis, pleometrosis independent colony founding Seifert and Schultz 2009; Seifert 2018 F0069 Formica dakotensis dakotensis free-living, TSP monogynous, polygynous monodomous, polydomous TSP, DCF: budding facultative temporary parasitism, high frequency budding Wasmann 1902, Talbot 1972 F0070 Formica dakotensis dakotensis free-living, TSP monogynous, polygynous monodomous, polydomous TSP, DCF: budding facultative temporary parasitism, high frequency budding Wasmann 1902, Talbot 1972 F0071 Formica dakotensis dakotensis free-living, TSP monogynous, polygynous monodomous, polydomous TSP, DCF: budding facultative temporary parasitism, high frequency budding Wasmann 1902, Talbot 1972 F0072 Formica densiventris difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding Mackay and Mackay 2002; SP Cover and C Rabeling unpubl. collection data F0073 Formica densiventris difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding Mackay and Mackay 2002; SP Cover and C Rabeling unpubl. collection data F0075 Formica cf densiventris difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover and C Rabeling unpubl. collection data F0164 Formica cf densiventris difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover and C Rabeling unpubl. collection data F0061 Formica dolosa pallidevulva free-living monogynous, polygynous monodomous ICF: haplometrosis, pleometrosis independent colony founding Trager et al. 2007, SP Cover unpubl. collection data F0063 Formica dolosa pallidevulva free-living monogynous, polygynous monodomous ICF: haplometrosis, pleometrosis independent colony founding Trager et al. 2007, SP Cover unpubl. collection data F0050 Formica emeryi sanguinea TSP, dulosis unknown monodomous TSP obligate temporary parasitism without budding (dulosis) Wheeler 1913; SP Cover unpubl. collection data F0034 Formica exsecta exsecta free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Skibińska 1982; Pisarski 1982; Brown and Keller 2000; Haag-Liautard et al. 2008; Goryunov 2015 F0030 Formica exsectoides exsecta free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Frumhoff and Ward 1992; SP Cover unpubl. collection data F0031 Formica exsectoides exsecta free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Frumhoff and Ward 1992; SP Cover unpubl. collection data F0032 Formica exsectoides exsecta free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Frumhoff and Ward 1992; SP Cover unpubl. collection data F0103 Formica foreliana “fusca” free-living monogynous, polygynous monodomous ICF: haplometrosis independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0003 Formica fossaceps integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP, DCF: budding facultative temporary parasitism, high frequency budding Buren 1942; SP Cover unpubl. collection data F0004 Formica cf fossaceps integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP, DCF: budding facultative temporary parasitism, high frequency budding SP Cover unpubl. collection data F0017 Formica cf frontalis rufa free-living, TSP unknown unknown unknown unknown F0115 Formica fuscocinerea “fusca” free-living monogynous, polygynous monodomous, polydomous, supercolonial ICF: haplometrosis, DCF: budding independent colony founding Seifert 2002; Seifert 2018 F0120 Formica gagates gagates free-living monogynous, polygynous monodomous, polydomous, supercolonial ICF: haplometrosis, claustral independent colony founding Bernard 1967; Seifert 2018 F0121 Formica glacialis “fusca” free-living monogynous, polygynous monodomous, polydomous ICF, DCF: budding independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0105 Formica gnava “fusca” free-living monogynous, polygynous monodomous ICF: haplometrosis independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0147 Formica gnava “fusca” free-living monogynous, polygynous monodomous ICF: haplometrosis independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0113 Formica hewitti “fusca” free-living monogynous, polygynous monodomous ICF independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0059 Formica incerta pallidevulva free-living monogynous, polygynous monodomous ICF: haplometrosis, pleometrosis independent colony founding Trager et al. 2007, SP Cover unpubl. collection data F0065 Formica incerta pallidevulva free-living monogynous, polygynous monodomous ICF: haplometrosis, pleometrosis independent colony founding Trager et al. 2007, SP Cover unpubl. collection data F0010 Formica cf integra integra free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding Creighton 1940; SP Cover unpubl. collection data F0005 Formica integroides integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP facultative temporary parasitism, low frequency budding Creighton 1940; SP Cover unpubl. collection data F0114 Formica integroides integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP facultative temporary parasitism, low frequency budding Creighton 1940; SP Cover unpubl. collection data F0015 Formica knighti difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding Buren 1944; SP Cover and C Rabeling unpubl. collection data F0036 Formica lasioides neogagates free-living monogynous, polygynous monodomous ICF: haplometrosis, claustral independent colony founding SP Cover unpubl. collection data F0040 Formica lasioides neogagates free-living monogynous, polygynous monodomous ICF: haplometrosis, claustral independent colony founding SP Cover unpubl. collection data F0130 Formica lemani “fusca” free-living monogynous, polygynous monodomous, polydomous ICF: haplometrosis, claustral independent colony founding Seppä et al. 2009; Gardner et al. 2007; Seifert 2018 F0021 Formica lugubris rufa free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Pamilo et al. 1994; Maeder and Cherix 2001; Gyllenstrand and Seppä 2003 F0037 Formica manni neogagates free-living monogynous monodomous ICF independent colony founding Wheeler 1913; Creighton 1950; JT Longino and PS Ward unpubl. collection data F0035 Formica mesasiatica exsecta free-living, TSP unknown monodomous, polydomous TSP, DCF: budding facultative temporary parasitism, high frequency budding Seifert 2000 F0129 Formica moki “fusca” free-living monogynous, polygynous monodomous ICF independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0117 Formica montana nr. “fusca” free-living monogynous, polygynous monodomous, polydomous ICF: haplometrosis, pleometrosis, DCF: budding independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0106 Formica neoclara “fusca” free-living monogynous, polygynous monodomous ICF independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0038 Formica neogagates neogagates free-living monogynous, polygynous monodomous ICF: haplometrosis, claustral independent colony founding Creighton 1940; Frumhoff and Ward 1992; SP Cover unpubl. collection data F0107 Formica neorufibarbis “fusca” free-living monogynous, polygynous monodomous ICF independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0109 Formica neorufibarbis “fusca” free-living monogynous, polygynous monodomous ICF independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0127 Formica neorufibarbis “fusca” free-living monogynous, polygynous monodomous ICF independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0128 Formica neorufibarbis “fusca” free-living monogynous, polygynous monodomous ICF independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0108 Formica neorufibarbis “fusca” free-living monogynous, polygynous monodomous ICF independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0126 Formica neorufibarbis “fusca” free-living monogynous, polygynous monodomous ICF independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0077 Formica nepticula difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding Wheeler 1913; SP Cover unpubl. collection data F0078 Formica nevadensis difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding Wheeler 1904; SP Cover unpubl. collection data F0079 Formica nevadensis difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding Wheeler 1904; SP Cover unpubl. collection data F0080 Formica nevadensis difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding Wheeler 1904; SP Cover unpubl. collection data F0007 Formica obscuripes integra free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Frumhoff and Ward 1992; Talbot 2012 F0008 Formica obscuripes integra free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Frumhoff and Ward 1992; Talbot 2012 F0009 Formica obscuripes integra free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Frumhoff and Ward 1992; Talbot 2012 F0162 Formica obscuripes integra free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Frumhoff and Ward 1992; Talbot 2012 F0006 Formica obscuriventris integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP facultative temporary parasitism, low frequency budding Creighton 1940; Frumhoff and Ward 1992; SP Cover and C Rabeling unpubl. collection data F0011 Formica obscuriventris integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP facultative temporary parasitism, low frequency budding Creighton 1940; Frumhoff and Ward 1992; SP Cover and C Rabeling unpubl. collection data F0046 Formica obtusopilosa neogagates free-living monogynous, polygynous monodomous ICF: haplometrosis, claustral independent colony founding Creighton 1950; SP Cover unpubl. collection data F0047 Formica obtusopilosa neogagates free-living monogynous, polygynous monodomous ICF: haplometrosis, claustral independent colony founding Creighton 1950; SP Cover unpubl. collection data F0131 Formica occulta nr. “fusca” free-living monogynous, polygynous monodomous, polydomous ICF: haplometrosis independent colony founding Francoeur 1973; Frumhoff and Ward 1992; SP Cover unpubl. collection data F0134 Formica orangea “fusca” free-living unknown unknown unknown unknown Seifert and Schultz 2009 F0012 Formica oreas integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP facultative temporary parasitism, low frequency budding Sather 1972; SP Cover and ML Borowiec unpubl. collection data F0013 Formica oreas integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP facultative temporary parasitism, low frequency budding Sather 1972; SP Cover and ML Borowiec unpubl. collection data F0125 Formica oreas integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP facultative temporary parasitism, low frequency budding Sather 1972; SP Cover and ML Borowiec unpubl. collection data F0135 Formica pacifica “fusca” free-living unknown unknown unknown unknown Francoeur 1973; JT Longino unpubl. collection data F0060 Formica pallidefulva pallidevulva free-living monogynous, polygynous monodomous ICF: haplometrosis, pleometrosis independent colony founding Trager et al. 2007; SP Cover unpubl. collection data F0066 Formica pallidefulva pallidevulva free-living monogynous, polygynous monodomous ICF: haplometrosis, pleometrosis independent colony founding Trager et al. 2007; SP Cover unpubl. collection data F0053 Formica pergandei sanguinea TSP, obl. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Trager et al. 2007; SP Cover unpubl. collection data F0051 Formica nr pergandei sanguinea TSP, obl. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Trager et al. 2007; SP Cover unpubl. collection data F0039 Formica perpilosa neogagates free-living monogynous, polygynous monodomous ICF: haplometrosis, claustral independent colony founding Wheeler 1913; SP Cover and C Rabeling unpubl. collection data F0028 Formica cf planipilis integra free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Creighton 1940; SP Cover unpubl. collection data F0145 Formica podzolica “fusca” free-living monogynous, polygynous monodomous, polydomous ICF: haplometrosis, DCF: budding independent colony founding Francoeur 1973; Frumhoff and Ward 1992; SP Cover unpubl. collection data F0019 Formica polyctena rufa free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Collingwood 1979; Ito and Higashi 1990; Seifert 2018 F0020 Formica pratensis rufa free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Horstmann et al. 1994; Sorensen and Schmidt 1983; Seifert 2018 F0033 Formica pressilabris exsecta free-living, TSP monogynous, polygynous monodomous, polydomous TSP, DCF: budding facultative temporary parasitism, high frequency budding Pamilo and Rosengren 1984; Seifert 2018 F0116 Formica propatula “fusca” free-living unknown unknown unknown unknown Francoeur 1973 F0016 Formica propinqua integra free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding Creighton 1940; SP Cover unpubl. collection data F0056 Formica puberula sanguinea TSP, obl. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Creighton 1950; SP Cover unpubl. collection data F0122 Formica puberula sanguinea TSP, obl. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Creighton 1950; SP Cover unpubl. collection data F0081 Formica querquetulana difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding Kennedy and Dennis 1937; SP Cover unpubl. collection data F0095 Formica querquetulana difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding Kennedy and Dennis 1937; SP Cover unpubl. collection data F0026 Formica ravida integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP, DCF: budding facultative temporary parasitism, high frequency budding Creighton 1940; SP Cover unpubl. collection data F0027 Formica ravida integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP, DCF: budding facultative temporary parasitism, high frequency budding Creighton 1940; SP Cover unpubl. collection data F0086 Formica reflexa dakotensis TSP with obligate dependence on monogynous monodomous TSP, perhaps obligatory permanent social parasitism Buren 1942, SP Cover unpubl. collection data host workers; possible inquilinism. Only mixed colonies; no evidence of dulosis F0054 Formica rubicunda sanguinea free-living, TSP, obl. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Creighton 1950; SP Cover unpubl. collection data F0055 Formica rubicunda sanguinea free-living, TSP, obl. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Creighton 1950; SP Cover unpubl. collection data F0018 Formica rufa rufa free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Collingwood 1979; Ito and Higashi 1990; Gösswald 2012; Seifert 2018 F0136 Formica rufibarbis “fusca” free-living monogynous, polygynous monodomous ICF: haplometrosis, pleometrosis independent colony founding Gammans 2008; Seifert and Schultz 2009 F0048 Formica sanguinea sanguinea free-living, TSP, fac. dulosis monogynous, polygynous monodomous, polydomous TSP, DCF: budding obligate temporary parasitism without budding (dulosis) Collingwood 1979; Frumhoff and Ward 1992; Mori and Le Moli 1998; Seifert 2018 F0049 Formica sanguinea sanguinea free-living, TSP, fac. dulosis monogynous, polygynous monodomous, polydomous TSP, DCF: budding obligate temporary parasitism without budding (dulosis) Collingwood 1979; Frumhoff and Ward 1992; Mori and Le Moli 1998; Seifert 2018 F0132 Formica sibylla “fusca” free-living unknown monodomous unknown unknown Francoeur 1973; PS Ward unpubl. collection data F0133 Formica cf sibylla “fusca” free-living unknown unknown unknown unknown PS Ward unpubl. collection data F0101 Formica sp. “fusca grp” “fusca” free-living unknown unknown unknown unknown SP Cover unpubl. collection data F0142 Formica sp. “fusca grp” “fusca” free-living unknown unknown unknown unknown SP Cover unpubl. collection data F0143 Formica sp. “fusca grp” “fusca” free-living unknown unknown unknown unknown SP Cover unpubl. collection data F0160 Formica sp. “fusca grp” “fusca” free-living unknown unknown unknown unknown SP Cover unpubl. collection data F0161 Formica sp. “fusca grp” “fusca” free-living unknown unknown unknown unknown SP Cover unpubl. collection data F0165 Formica sp. “fusca grp” “fusca” free-living unknown unknown unknown unknown SP Cover unpubl. collection data F0074 Formica sp. “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover unpubl. collection data F0087 Formica sp. “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover unpubl. collection data F0089 Formica sp. “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover unpubl. collection data F0090 Formica sp. “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover unpubl. collection data F0091 Formica sp. “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover unpubl. collection data F0092 Formica sp. “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover unpubl. collection data F0093 Formica sp. “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover unpubl. collection data F0094 Formica sp. “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover unpubl. collection data F0096 Formica sp. “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover unpubl. collection data F0157 Formica sp. “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding SP Cover unpubl. collection data F0058 Formica sp. “neogagates grp” neogagates free-living monogynous, polygynous monodomous ICF: haplometrosis, claustral independent colony founding SP Cover unpubl. collection data F0082 Formica sp. mlb01 “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding ML Borowiec unpubl. collection data F0083 Formica sp. mlb02 “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding ML Borowiec unpubl. collection data F0084 Formica sp. mlb03 “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding ML Borowiec unpubl. collection data F0085 Formica sp. mlb04 “micro grp” difficilis free-living, TSP monogynous, polygynous monodomous TSP facultative temporary parasitism, low frequency budding ML Borowiec unpubl. collection data F0123 Formica sp. sanguinea grp sanguinea TSP, dulosis (fac. or obl.?) unknown unknown unknown unknown SP Cover unpubl. collection data F0104 Formica subaenescens “fusca” free-living monogynous, polygynous monodomous, polydomous ICF: haplometrosis, pleometrosis independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0022 Formica subnitens integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP facultative temporary parasitism, low frequency budding Creighton 1940; SP Cover unpubl. collection data F0025 Formica subnitens integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP facultative temporary parasitism, low frequency budding Creighton 1940; SP Cover unpubl. collection data F0014 Formica cf subnitens integra free-living, TSP monogynous, polygynous monodomous, polydomous TSP facultative temporary parasitism, low frequency budding Creighton 1940; SP Cover unpubl. collection data F0137 Formica subpolita “fusca” free-living monogynous, polygynous monodomous, polydomous ICF: haplometrosis independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0138 Formica subpolita “fusca” free-living monogynous, polygynous monodomous, polydomous ICF: haplometrosis independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0144 Formica subsericea “fusca” free-living monogynous, polygynous monodomous, polydomous ICF: haplometrosis, pleometrosis independent colony founding Francoeur 1973; Frumhoff and Ward 1992; SP Cover unpubl. collection data F0146 Formica subsericea “fusca” free-living monogynous, polygynous monodomous, polydomous ICF: haplometrosis, pleometrosis independent colony founding Francoeur 1973; Frumhoff and Ward 1992; SP Cover unpubl. collection data F0159 Formica subsericea “fusca” free-living monogynous, polygynous monodomous, polydomous ICF: haplometrosis, pleometrosis independent colony founding Francoeur 1973; Frumhoff and Ward 1992; SP Cover unpubl. collection data F0156 Formica talbotae difficilis workerless inquilinism polygynous monodomous TSP permanent social parasitism Talbot 1977, Wilson 1977, Talbot 2012 F0141 Formica transmontanis “fusca” free-living unknown monodomous unknown independent colony founding Francoeur 1973; PS Ward unpubl. collection data F0023 Formica truncorum rufa free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Collingwood 1979; Frumhoff and Ward, 1992; Seifert 2018 F0029 Formica ulkei exsecta free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Holmquist 1928; SP Cover unpubl. collection data F0024 Formica uralensis uralensis free-living, TSP monogynous, polygynous monodomous, polydomous, supercolonial TSP, DCF: budding facultative temporary parasitism, high frequency budding Busch 1998; Münch 1991; Preuß 1956; Preuß 1957; Seifert 2018 F0057 Formica wheeleri sanguinea TSP, obl. dulosis monogynous, polygynous monodomous TSP obligate temporary parasitism without budding (dulosis) Creighton 1950; SP Cover unpubl. collection data F0139 Formica xerophila “fusca” free-living monogynous monodomous ICF: haplometrosis independent colony founding Francoeur 1973; SP Cover unpubl. collection data F0140 Formica cf xerophila “fusca” free-living monogynous monodomous ICF: haplometrosis independent colony founding PS Ward unpubl. collection data D1135 Iberoformica subrufa n/a free-living monogynous monodomous ICF independent colony founding Tinaut 1990; Muñoz-López et al. 2012; Seifert 2018 F0158 Polyergus longicornis n/a TSP, obl. dulosis monogynous monodomous TSP obligate temporary parasitism without budding (dulosis) Trager 2013 F0153 Polyergus mexicanus n/a TSP, obl. dulosis monogynous monodomous TSP obligate temporary parasitism without budding (dulosis) Trager 2013 F0154 Polyergus mexicanus n/a TSP, obl. dulosis monogynous monodomous TSP obligate temporary parasitism without budding (dulosis) Trager 2013 F0149 Polyergus rufescens n/a TSP, obl. dulosis monogynous monodomous TSP obligate temporary parasitism without budding (dulosis) Trager 2013 F0124 Polyergus vinosus n/a TSP, obl. dulosis monogynous monodomous TSP obligate temporary parasitism without budding (dulosis) Trager 2013 F0150 Proformica mongolica n/a free-living monogynous, polygynous monodomous ICF independent colony founding Galkowski et al. 2017 F0152 Rossomyrmex minuchae n/a TSP, obl. dulosis monogynous monodomous TSP obligate temporary parasitism without budding (dulosis) Ruano and Tinaut 2005

18 Supplementary references

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