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FIG. 15. Prothylacynus patagonicus MACN-A 5931 in Caudal Cranium Of Thylacosmilus Atrox (Mammalia, Metatheria, Sparassodonta), A South American Predaceous Sabertooth

FIG. 15. Prothylacynus patagonicus MACN-A 5931 from Corriguen-Kaik, Santa Cruz, Argentina; Santa Cruz Formation; Santacrucian, Early Miocene. Ventral view of A, the caudal cranium, with B, key. Abbreviations: AL, alisphenoid; bjs, basijugular sulcus; BO, basioccipital; BS, basisphenoid; cc, carotid canal; cod, occipital condyle; codf, condyloid foramen; cs, carotid sulcus; eam, external acoustic meatus; ewsq, squamosal epitympanic wing; EX, exoccipital; fmg, foramen magnum; fo, foramen ovale; fs, facial sulcus; hf, hypoglossal foramen; ips, inferior petrosal sinus; jf, jugular foramen; pcp, paracondylar process; PE, petrosal; pf, piriform fenestra; pgf, postglenoid foramen; ptp, posttympanic process; SQ, squamosal; tc, transverse canal. In this taxon, the more rostral foramen, opening into the basijugular sulcus, is tiny, and it presumably did not carry the hypoglossal nerve. For this reason, we identify it as a strictly venous foramen. Asterisk (*) indicates putative suture between alisphenoid and squamosal. Arrow indicates the direction to or the position of features hidden by other structures.

FIG. 13 in Caudal Cranium Of Thylacosmilus Atrox (Mammalia, Metatheria, Sparassodonta), A South American Predaceous Sabertooth

FIG. 13. Cladosictis patagonica YPM PU 15170, from locality 15 km south of Coy Inlet, Santa Cruz, Argentina; Santa Cruz Formation, Santacrucian, Early Miocene. Stereo pairs of left ectotympanic in A, lateral and B, medial views, with C, D, keys. Abbreviations: cac, caudal crus; chtym, groove for chorda tympani nerve (CNVII); cty, crista tympani; gma, groove for rostral process of malleus; roc, rostral crus; stp, styliform process (= anterolateral process of the ectotympanic); tyi, tympanic incisure. Asterisk (*) indicates tubercles.

FIG. 19 in Caudal Cranium Of Thylacosmilus Atrox (Mammalia, Metatheria, Sparassodonta), A South American Predaceous Sabertooth

FIG. 19. Thylacosmilus atrox, paratype, FMNH P14344. Model of the left petrosal based on 3-D reconstruction from micro-CT data, in A, tympanic, B, lateral, C, and cerebellar views. The petrosal is damaged and the osseous labyrinth is partly exposed. Abbreviations: amf, anteromedial flange; asc, anterior semicircular canal (exposed by breakage); chlc, cochlear canaliculus (= aqueductus cochlearis); co, cochlea (exposed by breakage); csc, cavum supracochleare (exposed by breakage); ctpp, caudal tympanic process of petrosal; ffc, fossula fenestrae cochleae (= round window); fsm, fossa for stapedius muscle; fv, fenestra vestibuli (= oval window); gpn, groove for greater petrosal nerve; iam, internal acoustic meatus; ips, sulcus for inferior petrosal sinus; lsc, lateral semicircular canal (exposed by breakage); pr, promontorium; psc, posterior semicircular canal (exposed by breakage); saf, subarcuate fossa; sff, secondary facial foramen; tt, tegmen tympani; ttf, tensor tympani fossa. Asterisk (*) indicates breakage.

FIG. 24 in Caudal Cranium Of Thylacosmilus Atrox (Mammalia, Metatheria, Sparassodonta), A South American Predaceous Sabertooth

FIG. 24. Monodelphis domestica (DUCEC), pouch young, coronal sections through the auditory region. In A–C, the internal carotid artery is seen passing into the endocranium in company with the internal carotid vein. Also indicated is the formation of the nerve of the pterygoid canal by the conjunction of the greater petrosal and deep petrosal nerves. In D, note position of the inferior petrosal sinus, situated intracranially adjacent to the central stem and otic capsule. Abbreviations: AL, alisphenoid; at, auditory tube; BS, basisphenoid; bsp, basilar plexus; cars, cavernous sinus; CNV, trigeminal (cranial) nerve; CNV3, mandibular branch of trigeminal (cranial) nerve; co, cochlear duct; ct, tympanic cavity; D, dentary; dpn, deep petrosal nerve (icn fibers passing out of tympanic plexus); EC, ectotympanic; fm, fibrous membrane of tympanic cavity; G, gonial; gg, geniculate ganglion (CNVII); gpn, greater petrosal nerve (CNVII); hy, hypophysis; ica, internal carotid artery; icn, internal carotid nerve (to tympanic plexus); icv, internal carotid vein; ips, inferior petrosal sinus; lpn, lesser petrosal nerve (CNIX); Mc, Meckel's cartilage; mpt, medial pterygoid muscle; mxv, maxillary vein; nph, nasopharynx; oc, oral cavity; ot, otic capsule; pa, processus alaris; phv, pharyngeal venous plexus; ttm, tensor tympani muscle. Histological serial section number indicated at top left. Numbers ascend in caudal direction.

FIG. 23 in Caudal Cranium Of Thylacosmilus Atrox (Mammalia, Metatheria, Sparassodonta), A South American Predaceous Sabertooth

FIG. 23. Thylacosmilus atrox, paratype, FMNH P14344. Pneumatization of the left middle ear based on 3-D reconstructions from micro-CT data in A, lateral, B, medial, and C, rostral views. Osseous labyrinth in pink and petrosal shown as semitransparent. Minor evaginations of the ventral paratympanic space not separately identified. Abbreviations: cps, caudal paratympanic space; ct, tympanic cavity; eam, external acoustic meatus; lps, lateral paratympanic space (= epitympanic sinus); vps, ventral paratympanic space.

FIG. 18. Sarcophilus lanarius AMNH M 65673 in Caudal Cranium Of Thylacosmilus Atrox (Mammalia, Metatheria, Sparassodonta), A South American Predaceous Sabertooth

FIG. 18. Sarcophilus lanarius AMNH M 65673. Caudal cranium in A, ventral and B, detailed oblique views. The tympanic floor is formed by the ectotympanic in the meatal area, plus the alisphenoid tympanic process and the rostral and caudal tympanic processes of the petrosal (last two fused into one structure). The exoccipital makes no contribution. Note extratympanic piriform fenestra. Abbreviations: AL, alisphenoid; at, aperture for auditory tube; bjs, basijulugar sulcus; BO, basioccipital; BS, basisphenoid; cc, carotid canal; cod, occipital condyle; ctpp, caudal tympanic process of petrosal; eam, external acoustic meatus; EC, ectotympanic; ewal, alisphenoid epitympanic wing; EX, exoccipital; fmg, foramen magnum; fo, foramen ovale (represented by its secondary aperture, which is bridged by a projection of the alisphenoid tympanic process—the anteromedial bullar lamina, following Pavan and Voss, 2016); gf, glenoid fossa; hfc, caudal hypoglossal foramen; hfr, rostral hypoglossal foramen; ips, inferior petrosal sinus; jf, jugular foramen; lps, lateral paratympanic space (= epitympanic sinus); pcp, paracondylar process; PE, petrosal; pf, piriform fenestra; pgf, postglenoid foramen; pgp, postglenoid process; pr, promontorium; ptp, posttympanic process; rps, rostral paratympanic space; rtpp, rostral tympanic process of petrosal; smf, stylomastoid foramen; SQ, squamosal; tc, transverse canal; tpal, tympanic process of alisphenoid. Arrow indicates the direction to or the position of features hidden by other structures.

FIG. 22 in Caudal Cranium Of Thylacosmilus Atrox (Mammalia, Metatheria, Sparassodonta), A South American Predaceous Sabertooth

FIG. 22. Thylacosmilus atrox, paratype, FMNH P14344, showing the angular relationship between plane of horizontal lateral semicircular canal (LSC) and the floor of the skull following De Beer (1947). A, Reconstruction of skull in lateral view oriented with horizontal LSC; B, inner ear in lateral view as positioned in the head; and C, parasagittal section, illustrating method of measurement (see text).

FIG. 27 in Caudal Cranium Of Thylacosmilus Atrox (Mammalia, Metatheria, Sparassodonta), A South American Predaceous Sabertooth

FIG. 27. Developmental contrasts in ontogeny of carotid foramen in representative marsupial and placental taxa: A, Lepus sp. (reconstruction, 45 mm CRL); B, Perameles sp. (reconstruction, 23 mm CRL); C, Monodelphis domestica (ZIUT; 15d postpartum, head, cleared and stained); D, Macropus eugenii (29 mm CRL); and E, Hemicentetes semispinosus (MPIH 1964/45, near-term fetus; 40 mm CRL). Abbreviations for A and B (from De Beer, 1937): acc, alicochlear commissure; ali, alisphenoid bone; at, ala temporalis; bcf, basicochlear (= basicapsular) fissure; bo, basioccipital bone; bs, basisphenoid bone; ds, dorsum sellae; exo, exoccipital bone; fac, internal acoustic meatus; fc, carotid foramen; fen, endolymphatic canal; fo, optic foramen; fpc, foramen prechiasmaticum; fpo, foramen pseudopticum; fr, foramen rotundum; fro, frontal bone; fsa, subarcuate fossa; hf, hypoglossal foramen; ip, interparietal; jug, jugal bone; lpc, lateral prefacial commissure; max, maxillary bone; nas, nasal bone; pal, processus alaris; par, parietal bone; pfc, prefacial commissure; pr, preoptic root of orbital cartilage; sob, supraoccipital bone; sq, squamosal bone. Abbreviations for C –E: AL, alisphenoid; at, auditory tube; BS, basisphenoid; cars, cavernous sinus; CNV, trigeminal (cranial) nerve; co, cochlear duct; ct, cavum tympani (= tympanic cavity); D, dentary; EC, ectotympanic; fc, carotid foramen; G, gonial; gpn, greater petrosal nerve (CNVII); ica, internal carotid artery; icn, internal carotid nerve; icv, internal carotid vein; Mc, Meckel's cartilage; mpt, medial pterygoid muscle; pa, processus alaris; pf, piriform fenestra; pr, promontorium; SQ, squamosal; tpbs, tympanic process of basisphenoid; ttm, tensor tympani muscle. In E, pair of asterisks (*) indicates the cartilaginous alichochlear commissures being replaced by bone, forming the definitive carotid foramen.

FIG. 14 in A sabre-tooth predator from the Neotropics: Cranial morphology of Anachlysictis gracilis Goin, 1997 (Metatheria, Thylacosmilidae), based on new specimens from La Venta (Middle Miocene, Colombia)

FIG. 14. — Bayesian majority-rule consensus topology of the post-burnin sample of trees for selected taxa (eutherian outgroup and metatherian ingroup), including the thylacosmilid sparassodont Anachlysictis gracilis Goin, 1997, focus of this study. Posterior probability support values> 75% indicated in branches.

FIG. 16 in A sabre-tooth predator from the Neotropics: Cranial morphology of Anachlysictis gracilis Goin, 1997 (Metatheria, Thylacosmilidae), based on new specimens from La Venta (Middle Miocene, Colombia)

FIG. 16. — Sequential reconstruction of the skull of Anachlysictis gracilis Goin, 1997: A, modified skull with original mandible; B, smoothed skull after mirroring; C, skull with reconstructed eyes, masticatory muscles and a tongue; D, 3D model of a reconstructed head. Artist: Tatsuya Shinmura. Scale bar: 20 mm.

FIG. 11 in A sabre-tooth predator from the Neotropics: Cranial morphology of Anachlysictis gracilis Goin, 1997 (Metatheria, Thylacosmilidae), based on new specimens from La Venta (Middle Miocene, Colombia)

FIG. 11. — Anachlysictis gracilis Goin, 1997; VPPLT 1612, lower dentition. Left lower dentition in occlusal (A), labial (E) and lingual (F) views; and right lower dentition in occlusal (B), labial (C) and lingual (D) views. Scale bar: 20 mm.

FIG. 8 in A sabre-tooth predator from the Neotropics: Cranial morphology of Anachlysictis gracilis Goin, 1997 (Metatheria, Thylacosmilidae), based on new specimens from La Venta (Middle Miocene, Colombia)

FIG. 8. — Anachlysictis gracilis Goin, 1997; VPPLT 1612, posterior portion of the skull: A, detail of basicranium in ventral view; B, lineal draw of the basicranium in ventral view; C, occipital view of the skull. Abbreviations: Al, alisphenoid; Bo, basioccipital; Bs, basisphenoid; bjs, basijugular sulcus; cf, carotid foramen; Eo, exoccipital; fm, foramen magnum; fo, foramen ovale; gf, glenoid fossa; hf, hypoglossal foramina; ips, foramen for the inferior petrosal sinus; jf, jugular fossa; Ju, jugal; pap, paracondylar process of exoccipital; pgp, postglenoid process; prgp, preglenoid process of jugal; Pt, pterygoid; spht, sphenoidal tubercles (basilar tubercles); Sq, squamosal. Scale bar: 20 mm.

FIG. 5 in A sabre-tooth predator from the Neotropics: Cranial morphology of Anachlysictis gracilis Goin, 1997 (Metatheria, Thylacosmilidae), based on new specimens from La Venta (Middle Miocene, Colombia)

FIG. 5. — Anachlysictis gracilis Goin, 1997; VPPLT 1612, skull in ventral view (A) and lineal draw (B). Abbreviations: Al, alisphenoid; Bo, basioccipital; Bs, basisphenoid; Eo, exoccipital; gf, glenoid fossa; hf, hypoglossal foramina; inf, incisive foramen; Ju, jugal; mpf, minor palatine foramen; mpp, medial palatine process of premaxilla; Mx, maxilla; n, notch; p, pits; pafo, paracanine fossa; Pal, palatine; pgp, postglenoid process; Psph, presphenoid; Pt, pterygoid; Px, premaxilla; Sq, squamosal. Scale bar: 20 mm.

FIG. 3 in A sabre-tooth predator from the Neotropics: Cranial morphology of Anachlysictis gracilis Goin, 1997 (Metatheria, Thylacosmilidae), based on new specimens from La Venta (Middle Miocene, Colombia)

FIG. 3. — Anachlysictis gracilis Goin, 1997; UCMP 39705, posterior fragment of right mandibular ramus, in lateral (A), posterior (B) and medial (C) views. Scale bar: 10 mm.

FIG. 4 in A sabre-tooth predator from the Neotropics: Cranial morphology of Anachlysictis gracilis Goin, 1997 (Metatheria, Thylacosmilidae), based on new specimens from La Venta (Middle Miocene, Colombia)

FIG. 4. — Anachlysictis gracilis Goin, 1997; VPPLT 1612, skull in dorsal view (A) and lineal draw (B). Abbreviations: Ip, interparietal; Ju, jugal; La, lacrimal; Mx, maxilla; Na, nasal; Pa, parietal; Pal, palatine; pop, postorbital process; Px, premaxilla; smf, suprameatal foramen; Sq, squamosal. Scale bar: 20 mm.

FIG. 1 in A sabre-tooth predator from the Neotropics: Cranial morphology of Anachlysictis gracilis Goin, 1997 (Metatheria, Thylacosmilidae), based on new specimens from La Venta (Middle Miocene, Colombia)

FIG. 1. — Geographical setting of the study area, indicating the collection points of specimens referred to Anachlysictis gracilis Goin, 1997 and generalized stratigraphic position. 1, IGM locality 75 (specimen IGM 184247, holotype); 2, UCMP locality V4531 ("Cerro Gordo 2": specimen UCMP 39705); 3, VPPLT locality "Finca Tres Pasos" (specimen VPPLT-1612).

FIG. 15. — 3D in A sabre-tooth predator from the Neotropics: Cranial morphology of Anachlysictis gracilis Goin, 1997 (Metatheria, Thylacosmilidae), based on new specimens from La Venta (Middle Miocene, Colombia)

FIG. 15. — 3D digital models of the skull of Anachlysictis gracilis Goin, 1997, based on the specimen VPPLT 1612. Original 3D scans of the skull in lateral (A) and anterior (B) views; modified retrodeformed skull with original unmodified mandible, in lateral (C) and anterior (D) views. Scale bar: 20 mm.

FIG. 13 in A sabre-tooth predator from the Neotropics: Cranial morphology of Anachlysictis gracilis Goin, 1997 (Metatheria, Thylacosmilidae), based on new specimens from La Venta (Middle Miocene, Colombia)

FIG. 13. — Consensus tree obtained from parsimony cladistic analysis under equal weights. Bremer supports (from 6411 trees, cut 0) indicated over branches; Bootstrap support values indicated below branches. Supports not given for Dasyuromorphia, as this node was constrained a priori (see text).

FIG. 10 in A sabre-tooth predator from the Neotropics: Cranial morphology of Anachlysictis gracilis Goin, 1997 (Metatheria, Thylacosmilidae), based on new specimens from La Venta (Middle Miocene, Colombia)

FIG. 10. — Anachlysictis gracilis Goin, 1997; VPPLT 1612, upper dentition. Right row in occlusal (A) and lingual (B) views; left side in occlusal (C) and lingual (D) views; and detail of the incisors area in occlusal view (E). Scale bars: 20 mm.