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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.

Figure 24 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 24. Photographs and line drawings of the hindlimb elements of Ligabuesaurus leanzai. A–E, right tibia MCF-PVPH-233/18 in anterior (A), lateral (B), posterior (C), proximal (D) and distal (E) views. F–I, proximal epiphysis of the left tibia MCF-PVPH-261/13 in proximal (F), posterior (G), lateral (H) and medial (I) views. J–M, right fibula MCF-PVPH-233/19 in medial (J), anterior (K), proximal (L) and distal (M) views. N, O, proximal epiphysis of the left fibula MCF-PVPH-261/14 in proximal (N) and medial (O) views. Abbreviations: amc, anteromedial crest; amf, anteromedial fossa; ap, anterior process; asap, articular surface for the ascending process; asp, astragal process; at, anterior trochanter; cc, cnemial crest; cf, cnemial fossa; lt, lateral trochanter; mi, insertion surface for muscle iliofibularis; pp, posterior process; tas, tibial articular surface. Scale bars: 10 cm.

Figure 23 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 23. Photographs and line drawings of the femora of Ligabuesaurus leanzai. A–C, right femur MCF-PVPH-233/17 in anterior (A), posterior (B) and distal (C) views. D–G, left femur MCF-PVPH-261/12 in anterior (D), distal (E), posterior (F) and medial (G) views. Abbreviations: ec, epicondyle; fc, fibular condyle; fh, femoral head; ft, fourth trochanter; ftr, femoral trochlea; gt, great trochanter; icf, intracondylar fossa; icg, intracondylar groove; lb, lateral bulge; lic, linea intermuscularis cranialis; tc, tibial condyle. Scale bars: 10 cm.

Figure 22 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 22. Photographs and line drawings of pelvic girdle elements of Ligabuesaurus leanzai. A, left ilium MCF-PVPH-261/08 in lateral view. B–D, proximal half of the right pubis MCF-PVPH-261/10 in lateral (B), proximal (C) and medial (D) views. E, F, proximal half of the left pubis MCF-PVPH-261/09 in lateral (E) and proximal (F) views. G, reconstruction of right pubis in lateral view, assembling the proximal and distal halves MCF-PVPH-261/10 and MCF-PVPH-261/11. H–J, distal half of the right pubis MCF-PVPH-261/11 in anterior (H), lateral (I) and distal (J) views. Abbreviations: acet, acetabulum; am, anterior margin; ias, ischiatic articular surface; ip, ischiatic peduncle; lm, lateral margin; mm, medial margin; of, obturator foramen; poap, postacetabular process; pp, pubic peduncle; prap, preacetabular process. Scale bars: 10 cm.

Figure 20 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 20. Photographs and line drawings of the forelimb elements of Ligabuesaurus leanzai. A–D, left humerus MCF-PVPH-233/10 in anterior (A), proximal (B), distal (C) and posterior (D) views. E, F, distal half of the right humerus MCF-PVPH-233/12 in anterior (E) and distal (F) views. G–I, distal half of the left radius MCF-PVPH-261/07 in posterior (G), distal (H) and lateral (I) views. Abbreviations: ac, accessory condyles; anf, anconeal fossa; das, distal articular surface; dpc, deltopectoral crest; dpf, deltopectoral fossa; hh, humeral head; ltf, lateral triceps fossa; mtf, medial triceps fossa; rc, radial condyle; uas, ulnar articular surface; uc, ulnar condyle. Scale bars: 10 cm.

Figure 19 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 19. Pectoral girdle elements of Ligabuesaurus leanzai. A, Right scapula MCF-PVPH-228/11 in lateral view. B, left scapula MCF-PVPH-233/08 in lateral view. C–E, left coracoid MCF-PVPH-261/05 in lateral (C), distal (D) and medial views (E). F–H, right coracoid MCF-PVPH-261/06 in medial (F), distal (G) and lateral (H) views. Abbreviations: af, anterior fossa; ar, acromial ridge; br, attachment surface for muscle biceps brachii; cbb, attachment surface for muscle coracobrachialis brevis; f, coracoid foramen; gl, glenoid; ign, infraglenoid notch; lev sc, attachment surface for levator scapulae; mvp, medioventral process; pf, posterior fossa; scs, scapulocoracoid articular surface; sr, scapular ridge. Scale bars: 10 cm.

Figure 15 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 15. Photograph and line drawings of the mid-posterior dorsal vertebrae MCF-PVPH-233/04 and MCF-PVPH-233/05 of Ligabuesaurus leanzai in left lateral view. Abbreviations: acdl, anterior centrodiapophyseal lamina; acpl, anterior centroparapophyseal lamina; pa, parapophysis; pcdl, posterior centrodiapophyseal lamina; pcpl, posterior centroparapophyseal lamina; pl, pleurocoel; poz, postzygapophysis; ppdl, parapophysisdiapophyseal lamina; prz, prezygapophysis. Scale bar: 10 cm.

Figure 16 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 16. Photographs and line drawings of the mid-posterior dorsal vertebrae MCF-PVPH-228/03 and MCF-PVPH-228/04 of Ligabuesaurus leanzai in left lateral (A), anterior (B) and posterior (C) views. Abbreviations: acdl, anterior centrodiapophyseal lamina; acpl, anterior centroparapophyseal lamina; cpol, centropostzygapophyseal lamina; hypa, hypantrum; pcdl, posterior centrodiapophyseal lamina; pcpl, posterior centroparapophyseal lamina; pl, pleurocoel; poz, postzygapophysis; ppdl, parapophysisdiapophyseal lamina; prz, prezygapophysis; spdl, spinodiapophyseal lamina; spol, spinopostzygapophyseal lamina; sprl, spinoprezygapophyseal lamina. Scale bar: 10 cm.

Figure 13 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 13. Photographs and line drawings of the anterior dorsal vertebra MCF-PVPH-233/03 of Ligabuesaurus leanzai in anterior (A), posterior (B) and left lateral (C) views. Abbreviations: acdl, anterior centrodiapophyseal lamina; cpol, centropostzygapophyseal lamina; cprl, centroprezygapophyseal lamina; di, diapophysis; lat. sprl, lateral spinoprezygapophyseal lamina; med. sprl, medial spinoprezygapophyseal lamina; nc, neural canal; pa, parapophysis; pcdl, posterior centrodiapophyseal lamina; pl, pleurocoel; poz, postzygapophysis; prsl, prespinal lamina; prz, prezygapophysis; spdl, spinodiapophyseal lamina; spol, spinopostzygapophyseal lamina; tprl, intraprezygapophyseal lamina. Scale bar: 10 cm.

Figure 21 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 21. Photographs and line drawings of the manus of Ligabuesaurus leanzai. A–D, right metacarpal II MCF-PVPH-233/13 in medial (A), proximal (B), distal (C) and lateral (D) views. E, distal epiphysis of the left metacarpal II MCF-PVPH-233/15 in distal view. F–I, right metacarpal III MCF-PVPH-233/14 in lateral (F), proximal (G), distal (H) and medial (I) views. J, distal epiphysis of the left metacarpal IV MCF-PVPH-233/16 in distal view. Abbreviations: am, anterior margin; lm, lateral margin; McI, articular surface for metacarpal I; McII, articular surface for metacarpal II; McIII, articular surface for metacarpal III; McIV, articular surface for metacarpal IV; pm, posterior margin; vlr, ventrolateral ridge. Scale bars: 10 cm.

Figure 12 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 12. Photographs and line drawings of the posterior cervical vertebra MCF-PVPH-233/02 of Ligabuesaurus leanzai in anterior (A), posterior (B) and right lateral (C) views. Abbreviations: acdl, anterior centrodiapophyseal lamina; cpol, centropostzygapophyseal lamina; cprl, centroprezygapophyseal lamina; di, diapophysis; lat. sprl, lateral spinoprezygapophyseal lamina; med. sprl, medial spinoprezygapophyseal lamina; nc, neural canal; ns, neural spine; pa, parapophysis; pcdl, posterior centrodiapophyseal lamina; pl, pleurocoel; podl, postzygodiapophyseal lamina; posl, postspinal lamina; poz, postzygapophysis; prdl, prezygodiapophyseal lamina; prz, prezygapophysis; r?, fragment of rib; spdl, spinodiapophyseal lamina; spol, spinopostzygapophyseal lamina; sprl, spinoprezygapophyseal lamina. Scale bar: 10 cm.

Figure 25 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 25. Photographs and line drawings of the right astragalus MCF-PVPH-233/20 of Ligabuesaurus leanzai in anterior (A), dorsal (B), lateral (C), posterior (D), ventral (E) and medial (F) views. Abbreviations: apr, ascending process of astragalus; cas, calcaneal articular surface; cpf, crest of posterior fossa; das, distal articular surface; fas, fibular articular surface; ilg, intertarsal ligament groove; pf, posterior fossa; tas, tibial articular surface. Scale bar: 10 cm.

Figure 14 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 14. Photograph and line drawing of the anterior dorsal vertebra MCF-PVPH-908 of Ligabuesaurus leanzai in anterior view. Abbreviations: acdl, anterior centrodiapophyseal lamina; cprl, centroprezygapophyseal lamina; di, diapophysis; lat. sprl, lateral spinoprezygapophyseal lamina; med. sprl, medial spinoprezygapophyseal lamina; pa, parapophysis; prsl, prespinal lamina; prz, prezygapophysis; spdl, spinodiapophyseal lamina; sprf, spinoprezygapophyseal fossa; tprl, intraprezygapophyseal lamina. Scale bar: 10 cm.

Figure 17 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 17. Photographs and line drawings of the posterior dorsal vertebrae MCF-PVPH-233/06 and MCF-PVPH-233/07 of Ligabuesaurus leanzai in anterior (A) and posterior (B) views. Abbreviations: acc. spdl, accessory spinodiapophyseal lamina; cpol, centropostzygapophyseal lamina; di, diapophysis; hypo, hyposphene; pcdl, posterior centrodiapophyseal lamina; poz, postzygapophysis; prsl, prespinal lamina; prz, prezygapophysis; spdl, spinodiapophyseal lamina; sprl, spinoprezygapophyseal lamina. Scale bar: 10 cm.

Figure 18 in Osteology and phylogenetic relationships of Ligabuesaurus leanzai (Dinosauria: Sauropoda) from the Early Cretaceous of the Neuquén Basin, Patagonia, Argentina

Figure 18. Photographs and line drawings of the caudal vertebra and dorsal rib of Ligabuesaurus leanzai. A–E, anterior caudal vertebra MCF-PVPH-261/15 in anterior (A), posterior (B), left lateral (C), dorsal (D) and ventral (E) views. F, G, dorsal rib MCF-PVPH-228/10 in medial view (F), with a detail of the internal pneumatic structure on the process of the capitulum (G). Abbreviations: aas, anterior articular surface; cap, capitulum; nc, neural canal; pas, posterior articular surface; rs, rib shaft; tp, transverse process; tub, tuberculum; vf, vascular foramen. Scale bars: 10 cm.