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Fig. 9. 3D in The First Reconstruction of the Head Anatomy of a Cretaceous Insect, †Gerontoformica gracilis (Hymenoptera: Formicidae), and the Early Evolution of

Fig. 9. 3D reconstructions of the heads of †Gerontoformica gracilis (on the left) and crown Formicidae (on the right) illustrating character transitions at the root of crown Formicidae, i.e., excluding †Gerontoformica. The relevant characters and states are marked with cyan outlines for plesiomorphies and magenta outlines for apomorphies; where applicable, physical directions of state changes are marked by arrows of the same colors. Complete illustrations of character states in all investigated taxa can be found in the character list at the end of this contribution. Char. 7: Ventral view on the head of †G. gracilis and Brachyponera luteipes, showing a medium versus a long postgenal bridge.Char. 36: Dorsal view on the posterior portion of the head of †G. gracilis and Brachyponera luteipes, showing presence versus absence of ocelli. Char. 42: Ventral view on the head of †G. gracilis and Brachyponera luteipes, showing a rounded versus projecting hypostomal tooth. Char. 46: Frontal view of the oral foramen of †G. gracilis and Brachyponera luteipes, showing a hypostomal corner situated far lateral of the tip of the hypostomal process versus a hypostomal corner aligned with the tip of the hypostomal triangular process. Char. 78: Frontal view of the scapus of †G. gracilis and Formica rufa showing a short scapus relative to the flagellum versus a long one. Only part of the antennal flagellum is shown, as this is not entirely imaged in the used µCT-scan data. Char. 86: Lateral view of the mandibular articulation of †G. gracilis and F. rufa showing the atala either broadly and flatly bulging or narrowly and highly bulging.Char. 94 and 97: Dorsal view of the mandible of †G. gracilis and F. rufa showing a narrow mandibular blade and gnathal edge without denticles proximad the subapical tooth versus a basally broadened mandibular blade with basal angle and denticles developed on the gnathal edge. Char. 108: Frontal section through the M. craniomandibularis internus (0md1) of †G. gracilis and F. rufa showing all fibers directly attaching to the main apodeme (magenta) versus some fibers attaching on thin cuticular fibrillae (cyan). Char. 143: Dorsal view of the prepharynx digestive tract of †G. gracilis and Wasmannia showing one bundle of the lateral portion of M. pharyngoepipharyngalis (0pe1l) versus two bundles.

Fig. 5. 3D in The First Reconstruction of the Head Anatomy of a Cretaceous Insect, †Gerontoformica gracilis (Hymenoptera: Formicidae), and the Early Evolution of

Fig. 5. 3D reconstructions of the head of †Gerontoformica gracilis specimen CASENT0741232, showing skeleton and musculature of maxilla and labium. (A, F, H, and K) Dorsal view.(B, E, I, and L) Ventral view. (C, G, and J) Sagittal view,in (C) the left instead of right side of the head is shown. (D) Frontal view. (M) Lateral view. (A–C and G) Maxillary musculature. (D–F) Maxilla. (H–J) Labial musculature and salivary duct. (K–M) Labium. Abbreviations: 0hy3, M. tentoriohypopharyngalis; 0hy12, M. hypopharyngosalivaris; 0la5, M. tentoriopraementalis; 0la11, M. praementoparaglossalis; 0la12, M. praementoglossalis; 0la14, M. praementopalpalis externus; 0mx1, M. craniocardinalis externus; 0mx3, M. tentoriocardinalis; 0mx4, M. tentoriostipitalis anterior; 0mx6, M. stipitolacinialis; 0mx7, M. stipitogalealis; 0mx8, M. stipitopalpalis externus; bpb, basiparaglossal brush; cd, cardo; ga, galea; gl, glossa; hy, distal hypopharynx; lb, labrum; lc, lacinia; msf, medial stipital flange; mxc, maxillary comb; pgl, paraglossa; plb, labial palp; plbp, process of the labial palp; pmd, premental ditch; pml, prementum lateral face; pmv, prementum ventral face; pmx, maxillary palp; psm, postmentum; st, stipes; svd, salivary duct. Symbols: blue line, premental ditch.

Fig. 2 in The First Reconstruction of the Head Anatomy of a Cretaceous Insect, †Gerontoformica gracilis (Hymenoptera: Formicidae), and the Early Evolution of

Fig. 2. Slices of the µ-CT dataset showing histological preservation in the head of †Gerontoformica gracilis specimen CASENT0741232. (A) Frontal section showing the cells of the mandibular gland. (B) Transverse section showing the optic neuropils of the brain. (C) Transverse section showing the prepharynx slightly posterad the antennal insertions. (D) Frontal section showing the mandibular adductor, arrows indicating the only possible cuticular filaments. (E) Longitudinal section showing antennal muscles, olfactory lobes, and part of the pharyngeal gland. (F) Overview of sagittal section. Abbreviations: 0bu5, M. tentoriobuccalis, only fragments preserved; 0md1, M. craniomandibularis internus; ada, adductor apodeme; ammc?, likely the antennal mechanosensory and motor center; anm, antennal musculature; ba, amber 'bubble' displacing tissue; bpg, basiparaglossal brush; br, brain; bt, buccal tube; la, lamina; lo, lobula; mb, mushroom body; me, medulla; nan, antennal nerve; glo, glomeruli of the olfactory lobe; mdg, mandibular gland; om, fold representing the oral arm; om, ommatidia of the compound eye; ph, pharynx; pph, prepharynx; sog, suboesophageal ganglion; tr, transverse prepharyngeal ridges. Colors: beige/brown, cuticular structures (ada & bpg); green, alimentary canal; orange, muscles; purple, glands; yellow, nervous system.

Fig. 4. 3D in The First Reconstruction of the Head Anatomy of a Cretaceous Insect, †Gerontoformica gracilis (Hymenoptera: Formicidae), and the Early Evolution of

Fig. 4. 3D reconstructions of the head of †Gerontoformica gracilis specimen CASENT0741232, showing skeleton and musculature of mandible and antenna. (A and D) Dorsal view. (B and E) Ventral view. (C and F) Sagittal view. (G) Outer view. (H) Inner view. (A–C) Antennal musculature. (D–F) Mandibular musculature and mandibular gland. (G and H) Mandible. Abbreviations: 0an1, M. tentorioscapalis anterior; 0an2, M. tentorioscapalis posterior; 0an3, M. tentorioscapalis lateralis; 0an4, M. tentorioscapalis medialis; 0an6, M. scapopedicellaris lateralis; 0an7, M. scapopedicellaris medialis; 0md1, M. craniomandibularis internus; 0md3, M. craniomandibularis externus; aba, abductor apodeme; ada, adductor apodeme; adc, adductor carina; al, atala; at, apical tooth; bm, basal margin; cdc, condylar carina; cdg, condylar groove; dta, dorsal tentorial arm; fl, fimbriate line; ims, inner mandibular surface; ll, lateral tentorial lamella; ma, mandalus; md, mandible; mda, mandibular acetabulum; mdc, mandibular condyle; mdg, mandibular gland; ml, medial tentorial lamella; omc, outer mandibular carina; oms, outer mandibular surface; pd, pedicel; sat, subapical tooth; sc, scapus. Symbols: blue line, basal curvature of pedicel.

Fig. 6. 3D in The First Reconstruction of the Head Anatomy of a Cretaceous Insect, †Gerontoformica gracilis (Hymenoptera: Formicidae), and the Early Evolution of

Fig. 6. 3D reconstructions of the head of †Gerontoformica gracilis specimen CASENT0741232, showing the digestive tract with its muscles and glands and the central nervous system. (A, E, and G) Dorsal view. (B, C, and H) Ventral view. (D and F) Sagittal view. (A, B, and D) Overview of digestive tract with muscles and glands and central nervous system. (C) Head capsule cut open to reveal origin sites of dorsal head muscles. (E and F) Prepharynx with oral arm folds. (G and H) Recognizable subregions of the brain. Abbreviations: 0ci1a, M. clypeopalatalis a; 0ci1b, M. clypeopalatalis b; 0bu1, M. clypeobuccalis; 0bu2, M. frontobuccalis anterior; 0hy1, M. frontooralis; 0pe1d, M. pharyngoepipharyngalis dorsal bundle; 0pe1l, M. pharyngoepipharyngalis lateral bundle; ammc?, likely the antennal mechanosensory and motor center; aot, anterior optic tubercle; br, brain; bt, buccal tube; cbl, central body lower unit; cbu, central body upper unit; ce, compound eye;ep, epipharynx;fp, food pellet;hy, distal hypopharynx;ibp, infrabuccal pouch;la, lamina;lb, labrum; lbm, labium;lo, lobula;me, medulla;mbpd, mushroom body peduncle; mbca, mushroom body calyx; nan, antennal nerve; no, noduli; oa, oral arm fold; ol, olfactory lobe; opl, optic lobe; ph, pharynx; phg, pharyngeal gland; pph, prepharynx; pphg, prepharyngeal gland; sog, suboesophageal ganglion. Symbols: dotted magenta line, folds presumably representing the oral arms.

Fig. 7 in The First Reconstruction of the Head Anatomy of a Cretaceous Insect, †Gerontoformica gracilis (Hymenoptera: Formicidae), and the Early Evolution of

Fig. 7. Character state transformations recovered via parsimony and Bayesian analyses. Only unequivocally recovered transformations are shown.Topology fixed after Branstetter et al. (2017) and Boudinot et al. (2022c). Transformations are marked as open circles (homoplastic features) or solid boxes (unambiguous features), with the character number above and the transformation states below. Characters that are used here for the first time are marked in green, and those used in other analyses previously are marked in light orange.The timeline at the bottom is not to scale and not based on our analyses, but the marked time points (145, 99, and 66.5 ma) are positioned roughly to reflect current hypotheses on splitting events, mainly based on the divergence time estimations of Boudinot et al. (2022c). We set the branch tip of †Gerontoformica at 99 ma as this is the likely age of the fossil we investigated here.

Fig. 3. 3D in The First Reconstruction of the Head Anatomy of a Cretaceous Insect, †Gerontoformica gracilis (Hymenoptera: Formicidae), and the Early Evolution of

Fig. 3. 3D reconstructions of the head of †Gerontoformica gracilis specimen CASENT0741232, showing external and internal skeletal elements. (A and D) Dorsal view. (B and E) Ventral view. (C) Lateral view. (Cʹ) Dorsolateral view of the torulus. (F) Sagittal view. (G) Frontal view of the oral foramen of the head capsule. (A–C) Complete head. (D–F) Internal skeleton. (H) Labrum outer surface. (I) Labrum inner surface. Abbreviations: alf, atalar fossa; ant, antennifer; ata, anterior tentorial arm; atp, anterior tentorial pit; bb, bulbus; bbn, bulbus neck; ce, compound eye; cl, clypeus; cli, clypeal inflection; clp, clypeal process; clc, clypeal chaetae; cor, circumocular ridge; dma, dorsal mandibular articulation; dta, dorsal tentorial arm; ess, epistomal sulcus; esr, epistomal ridge; fc, frontal carina; fr, frontal area; hc, cardinal condyle of hypostoma; hygl, lateral hypostomal groove; hygm, medial hypostomal groove; hyp, hypostomal triangular process; hysci, inner hypostomal carina; hyscor, oral hypostomal carina; hyscou, outer hypostomal carina; hyt, hypostomal corner; ll, lateral tentorial lamella; lbre, outer labral surface; lbri, inner labral surface; ltra, transverse row of labral setae; md, mandible; ml, medial tentorial lamella; oc, ocellus; oca, occipital carina; occ, occipital area; ocf, occipital foramen; pgb, postgenal bridge; pgr, postgenal ridge; plb, labial palp; pmx, maxillary palp; poc, postocciput; pta, posterior tentorial arm; sc, scapus; sfg, subforaminal groove; st, stipes; tb, tentorial bridge; tba, tentorial bridge anteriomedian process; to, torulus; toi, inner torular rim; vma, pleurostomal fossa. Symbols: blue line, epistomal sulcus.

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Figure 1 in Evidence for the evolution of eusociality in stem ants and a systematic revision of †Gerontoformica (Hymenoptera: Formicidae)

Figure 1. Overview of the Kachin amber piece and its syninclusions. A, the pupa of †G. gracilis (CASENT01741231). B, the adult of †G. gracilis (CASENT0741232). C, the holotype of †G. sternorhabda sp. nov. (CASENT0741233). D, the paratype of †G. sternorhabda sp. nov. (CASENT0741234). E, the entire amber piece (AMNH SY-23).

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Figure 5 in Evidence for the evolution of eusociality in stem ants and a systematic revision of †Gerontoformica (Hymenoptera: Formicidae)

Figure 5. Photomicrographic details of †G. sternorhabda sp. nov. Paratype specimen (UFV-LABECOL-009656). A, head in approximate facial view. B, mouthparts in anterolateral oblique view. C, head and mesosoma in lateral view. D, proximal segments of metathoracic leg in approximate anterior view. E, metasoma in approximate lateral view. F, petiole in lateral view. G, prothoracic tarsus, oblique posterior view. H, metatibial apex and metabasitarsal base in posterior view. I, propretarsus in oblique distal view. J, metapretarsus in posteroventral oblique view. Abbreviations: AI, propodeum; AII– AVII, second through seventh abdominal segments; ca, calcar; cav, velum (lamella) of calcar; ce, compound eye; cl, clypeus; cll, lateral lobe of clypeus; cns, cinctus of fourth abdominal sternum; cnt, cinctus of fourth abdominal tergum; fc, frontal carina; hetg, helcial tergite; msctl, mesoscutellar area (mesoscutellum); msctm, mesoscutal area (mesoscutum); msnttc, transverse

Fig. 1 in The First Reconstruction of the Head Anatomy of a Cretaceous Insect, †Gerontoformica gracilis (Hymenoptera: Formicidae), and the Early Evolution of

Fig. 1. Photographic images of the amber fossil. (A) Overview of the amber piece:the solid arrowheads indicate synincluded adult †Gerontoformica sternorhabda, the open arrowhead indicates synincluded †G. gracilis pupa, and the circle outlines focal †G. gracilis specimen CASENT0741232. (B) Overview of specimen CASENT0741232 in ventrolateral view. (C) Head of †G. gracilis in frontolateral view (oblique dorsal); the short blue arrow indicates the process on the first labial palpomere. (D) Head of †G. gracilis in frontolateral view (oblique ventral). (E) Head of †G. gracilis in posterolateral view. Abbreviations: bb, bulbus; cl, clypeus; clc, clypeal chaetae; ce, compound eye; ess, epistomal sulcus; fc, frontal carina; fla, antennal flagellum; ga, galea; md, mandible; plb, labial palp; pmx, maxillary palp; sc, scapus.The photographs in A, B and D were also used in Boudinot et al. 2022b.

Fig. 8. 3D in The First Reconstruction of the Head Anatomy of a Cretaceous Insect, †Gerontoformica gracilis (Hymenoptera: Formicidae), and the Early Evolution of

Fig. 8. 3D reconstructions of the heads of †Gerontoformica gracilis (on the right) and outgroup taxa (on the left) illustrating character transitions at the root of the total clade Formicidae.The relevant characters and states are marked with cyan outlines for plesiomorphies and magenta outlines for apomorphies; where applicable, physical directions of state changes are marked by arrows of the same colors. Complete illustrations of character states in all investigated taxa can be found in the character list at the end of this contribution. Char. 24: Frontal view on the head of Sceliphron caementarium and †G. gracilis, showing a flat versus bulging frontal region between the toruli. Char. 34: Lateral view of the head of Ampulex sp. and †G. gracilis showing large versus small compound eyes. Char. 57: Sagittal view of the tentorium of Ampulex sp. and †G. gracilis showing a long versus a short dorsal tentorial arm. Char. 73: Outer surface of the labrum of Ampulex sp. and †G. gracilis showing absence versus presence of a transverse line of setae.Char. 79: Frontal view of the scapopedicellar articulation of Ampulex sp. and †G. gracilis showing a basally straight and cylindrical versus a basally curved and flattened pedicel, in addition to an anteriorly deeply notched scapus tip. Char. 83: Lateral view of the mandibular articulation of S. caementarium and †G. gracilis showing a short versus elongated dorsal (secondary) mandibular articulation. Char. 145: Dorsal view of the cephalic digestive tract of †G. gracilis showing presence of the pharyngeal gland (absence in outgroups not shown).