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FIG. 4. Didelphis virginiana TMM M-2517 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals

FIG. 4. Didelphis virginiana TMM M-2517 (Didelphidae, Didelphimorphia), adult specimen, endocast reconstruction showing osteological features associated with pericarotid venous network and related vasculature (data source, table 2). In this and all other figures depicting endocasts, unless indicated otherwise red-colored structures are casts of carotid canal, and thus represent both internal carotid artery and internal carotid vein. Blue-colored structures mostly represent venous conduits. Isolated portions of certain trackways have been reconnected to restore continuity of ventral petrosal sinus, extracranial continuation of ventral petrosal sinus, and sigmoid sinus. Views: A, ventral; B, same, with transverse basisphenoid sinus (in green) superimposed; C, oblique right lateral; D, oblique dorsal closeup of transverse canal junction (simple pattern); E, oblique caudoventral surface of intact caudal cranium. In D, note craniopharyngeal canal and interstitial canaliculi communicating with transverse canal along different planes. Canals actually open into transverse basisphenoid sinus, although this is not obvious because of color coding. Gap (asterisk) locates positions of cavernous sinus and pituitary. In E, extracranial continuation of ventral petrosal sinus reconstructed and shown passing, suc-

FIG. 5. Didelphis virginiana TMM M-2517 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals

FIG. 5. Didelphis virginiana TMM M-2517 (Didelphidae, Didelphimorphia), adult caudal cranium, selected coronal (A–H) and parasagittal (I–K) segments (data source, table 2). In A and B, note junction of rostral branches of transverse canals, communicating with but not enveloped by transverse basicranial sinus. Boundary between transverse basisphenoid sinus and regular diploe is gradational. In B–D, craniopharyngeal canal, of uncertain function, connects transverse canal junction with hypophyseal infundibular sulcus (see also I). In E, true caudal branches of transverse canals are absent; only short interstitial canaliculi represented. In J and K, pneumatized areas (possible additional sites of hematopoiesis) also seen in basioccipital and presphenoid). Key: AS, alisphenoid; BO, basioccipital; BS, basisphenoid; cc, carotid canal; cpc, craniopharyngeal canal; cpf, craniopharyngeal foramen; crp, cribriform plate; ctbs, caudal portion of transverse basisphenoid sinus; el, ethmoid labyrinth; encf, endocranial carotid foramen; encg, endocranial carotid groove; excf, exocranial carotid foramen; fm, foramen magnum; FR, frontal; his, hypophyseal infundibular sulcus; hpf, hypophyseal fossa; isc, interstitial canaliculus; junc, junction of transverse canals; junc + rtbs, combined junction and rostral transverse basisphenoid sinus; le, lateral extension of transverse basisphenoid

FIG. 15 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals

FIG. 15. Dromiciops gliroides MACN Ma-23607 (Microbiotheriidae, Microbiotheria), near-adult specimen, endocast reconstruction showing osteological features associated with pericarotid venous network and related vasculature (data source, table 2; color key, fig. 4). Skull is slightly damaged (e.g., left carotid canal). Views: A, ventral; B, same, with transverse basisphenoid sinus superimposed; C, oblique right lateral; D, horizontal segment through mesocranium; and E, ventral surface, intact caudal cranium. In C, sulcus for sigmoid sinus shallow and discontinuous; if internal jugular vein functionally present in adult, vessel probably small. In D, fingerlike projection from carotid canal leads to caudal branch foramen; transverse canals are continuous with transverse basicranial sinus, and rostral branches run through large compound space (cf. fig. 17A). Key: aptc, anterior pterygoid canal (=?anterior pterygoid foramen of Beck et al., 2023: char. 50); astp, tympanic process of alisphenoid; ats, sulcus for auditory tube; BO, basioccipital; cbf, caudal branch foramen; cc, carotid canal; ccs, sulcus leading to carotid canal; cchc, caudal condylohypoglossal canal; co, cochlea; ctbs, caudal portion of transverse basicranial sinus; etbs, eminence of transverse basisphenoid sinus; evpf, extracranial continuation of ventral petrosal sinus; hpf, hypophyseal fossa; jf, jugular foramen; le, lateral extension of transverse basisphenoid sinus; onvs, sulcus for ophthalmic neurovascular array; PAS, parasphenoid; PE, petrosal; pglf, postglenoid foramen; ptpf, pterygopalatine fissure; rbtc, rostral branch of transverse canal; rchc, rostral condylohypoglossal foramen; rtbs, rostral portion of transverse basicranial sinus; smf, suprameatal foramen; sss, sulcus for sigmoid sinus; tbs, transverse basicranial sinus; tcf, transverse canal foramen; tmc, tympanic cavity; tpss, sulcus for temporal sinus; vpss, sulcus for ventral petrosal sinus.

FIG. 25. Vombatus ursinus TMM M-2953 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals

FIG. 25. Vombatus ursinus TMM M-2953 (Vombatidae, Diprotodontia), adult caudal cranium, selected coronal segments in rostrocaudal order (data source, table 2). In A–C, large transverse canals meet in midline junction (cf. Phascolarctos, Aplin, 1990: 255). In D–G, note large right and left conduits for caudal branches of transverse canal, traceable bilaterally from caudal branch foramen in endocranial carotid groove to terminus in transverse canal trunk. Key: AS, alisphenoid; BS, basisphenoid; cbs, sulcus for caudal branch, leading into caudal branch foramen; cbtc, canal for caudal branch of transverse canal; cc, carotid canal; cpc, craniopharyngeal canal; cpf, craniopharyngeal foramen; ctbs, caudal portion of transverse basisphenoid sinus; encf, endocranial carotid foramen; encg, endocranial carotid groove; excf, exocranial carotid foramen; hpf,

FIG. 38 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals

FIG. 38. Sipalocyon gracilis AMNH VP-9254 (Hathliacynidae, Sparassodonta), adult specimen, endocast reconstruction showing osteological features associated with pericarotid venous network and related vasculature (data source, table 2; color key, fig. 4). Views: A, ventral; B, same, with transverse basisphenoid sinus superimposed; C, oblique right lateral; D, closeup of endocranial surface of mesocranial region; and E, caudoventral surface, intact caudal cranium showing extracranial continuation of ventral petrosal sinus. In A–C, rostral portion of transverse basicranial sinus lacks identifiable intramural connections with carotid canals or cavernous sinus. Feature 1, identified by Archer (1976) as a possible transverse canal in Sipalocyon, is more likely a channel for nerve of pterygoid canal. Feature 2 is a sulcus of unknown function, unrelated to either nerve of pterygoid canal or typical transverse canal. Feature 3, small channel within basisphenoid sinus tissue, presumably venous, passes between bilateral carotid canals (double-headed arrow in D) caudal to hypophyseal fossa. Arrangement is different from that of CBTC of investigated marsupials. Feature 4, another small canaliculus, either interstitial or remnant of notochord canal, connects feature 3 with rest of sinus. Key: cc, carotid canal; cchc, caudal condylohypoglossal canal; ctbs, caudal portion of

FIG. 29. Thylacinus cynocephalus AMNH M-144316 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals

FIG. 29. Thylacinus cynocephalus AMNH M-144316 (Thylacinidae, Dasyuromorphia), adult specimen, coronally hemisected skull. Views: A, ventral; B, oblique ventrolateral; C, oblique dorsocaudal; and D, coronal section through mesocranium, caudal aspect, closeup of endocranial carotid grooves. Asterisks mark union of carotid and transverse canals in base of carotid grooves (see fig. 31). In this version of hybrid configuration, interpretation is that enlarged caudal branch vein arising from cavernous sinus shares carotid groove with internal carotid neurovascular bundle before separating and departing through transverse canal to join trunk. For this reason,

FIG. 8. A, Caluromys derbianus AMNH M-18910 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals

FIG. 8. A, Caluromys derbianus AMNH M-18910 (Didelphidae, Didelphimorphia), adult caudal cranium in oblique caudoventral view. B, Caluromys sp. AMNH M-184599 (Didelphidae, Didelphimorphia), adult caudal cranium in endocranial view. Although transverse canal foramen is usually scored as completely absent in the short-tailed opossum group, in A a tiny dimple (single asterisk) lateral to exocranial opening of right carotid canal may represent vestige of vein's trackway, assuming it differentiated but did not persist. Equivalent opening is not seen on left side or in B. There is no swelling or eminence for transverse canals on endocranial floor, consistent with rostral branch tubes not being present. Endocranial carotid grooves contain only one aperture, for internal carotid neurovascular bundle, indicating caudal branch of transverse canal vein is also absent in this taxon. Note well marked foramina for craniopharyngeal (feature 1) and notochord (feature 2) canals.

FIG. 23. Distoechurus pennatus AMNH M-105938 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals

FIG. 23. Distoechurus pennatus AMNH M-105938 (Acrobatidae, Diprotodontia), adult caudal cranium in A, ventral, and B, oblique caudoventral views. Two sulci, incompletely separated by a bridge (b) formed by a calcified ligament, converge on a single aperture identified as exocranial carotid foramen. As there is no separate transverse canal foramen, transverse canal vein is thought to share carotid canal with internal carotid neurovascular bundle (Aplin, 1990). Basijugular sulcus not prominent; jugular foramen and extracranial continuation of ventral petrosal sinus are separate, but open into common external aperture. Key: b, bridge (calcified?sphenopetrosal ligament); BO, basioccipital; BS, basisphenoid; cchf, caudal condylohypoglossal foramen; ccs, sulcus leading to carotid canal; cdf, condylar foramen additional to rchf and cchf; eam, external acoustic meatus; evpf, extracranial continuation of ventral petrosal sinus; evpf + jf, common external aperture for jugular foramen and extracranial continuation of ventral petrosal sinus (left side only); excf, exocranial carotid foramen; fm, foramen magnum; fo, foramen ovale; jf, jugular foramen; pf, piriform fenestra; pglf, postglenoid foramen; rchf, rostral condylohypoglossal foramen; sof, sphenoorbital fissure; tcvs, sulcus for transverse canal vein.

FIG. 21. Trichosurus vulpecula TMM M-849 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals

FIG. 21. Trichosurus vulpecula TMM M-849 (Phalangeridae, Diprotodontia), adult specimen, endocast reconstruction showing osteological features associated with pericarotid venous network and related vasculature (data source, table 2; color key, fig. 4). Views: A, ventral; B, same, with transverse basisphenoid sinus superimposed; C, oblique right lateral; D, closeup of mesocranial osteological features in T. vulpecula AMNH 48055; and E, oblique caudoventral surface, intact caudal cranium. In B and C, endocast of caudal branch foramen projecting from carotid groove (cf. Dromiciops, fig. 15A, D). In A and D, in addition to caudal branch foramen, note foramina (single asterisk, box) perforating endocranial floor, linking hypophyseal fossa to transverse canals by means of hypophyseal canaliculi (cf. fig. 22C, D). Separate craniopharyngeal foramen not identifiable. In E, note foramen (double asterisks) medial to position of foramen ovale (see text). Key: atcf, accessory transverse canal foramen; atcl (?rbtc), accessory transverse canal (?rostral branch of transverse canal); cbf, caudal branch foramen; cc, carotid canal; cchc, caudal condylohypoglossal canal; cchf, caudal condylohypoglossal canal; ccs, sulcus leading to carotid canal; ctbs, caudal portion of transverse basisphenoid sinus; encf, endocranial carotid canal; encg, endocranial carotid groove; etbs, eminence of transverse basisphenoid sinus; evpf, extracranial continuation of ventral petrosal sinus; fo, foramen ovale; jf, jugular foramen; le, lateral extension of transverse basisphenoid sinus; mdnc, canal for mandibular nerve; mxns, sulcus for maxillary nerve; onvs, sulcus for

FIG. 14 in New material of Incadelphys antiquus (Pucadelphyda, Metatheria, Mammalia) from the early Palaeocene of Bolivia reveals phylogenetic affinities with enigmatic North and South American metatherians

FIG. 14. — Phylogenetic relationships of Incadelphys among other metatherians:strict consensus tree (Consistency index [CI] = 0.401; Retention index [RI] = 0.609) of the two shortest trees resulting from the analysis with downweighted homoplastic characters (with Goloboff k = 3); black number above nodes indicate the minimum and maximum numbers of synapomorphies. The name of clade designated as "SAMI" is a working term composed with the initials of the four genera it includes: Szalinia, Aenigmadelphys, Marmosopsis, and Incadelphys.

FIG. 12 in New material of Incadelphys antiquus (Pucadelphyda, Metatheria, Mammalia) from the early Palaeocene of Bolivia reveals phylogenetic affinities with enigmatic North and South American metatherians

FIG. 12. — Aenigmadelphys archeri, SEM photos of casts: A, left M3 (holotype, OMNH 23328) in occlusal view; B, right m3 (OMNH 20531)in occlusal view; C, the same in labial view; D, the same in lingual view. Scale bar: 1 mm.

FIG. 11 in New material of Incadelphys antiquus (Pucadelphyda, Metatheria, Mammalia) from the early Palaeocene of Bolivia reveals phylogenetic affinities with enigmatic North and South American metatherians

FIG. 11. — Occlusal view of the M1 of some basal metatherians discussed in text showing the distolabial extension of the metastylar lobe: A, Aenigmadelphys (OMNH 22898), SEM photo of cast; B, Incadelphys (YPFB Pal 6251), SEM photo of cast; C, Incadelphys (MHNC 13906); D, Marmosopsis (MNHN.F.ITB83). Scale bar: 1 mm.

FIG. 10 in New material of Incadelphys antiquus (Pucadelphyda, Metatheria, Mammalia) from the early Palaeocene of Bolivia reveals phylogenetic affinities with enigmatic North and South American metatherians

FIG. 10. — Incadelphys antiquus, MHNC 13906: A, lateral view of the right dentary; B, medial view of the right dentary; C, lateral view of the left dentary; D, medial view of the left dentary; E, ventral view of the left dentary. Scale bar: 5 mm.

FIG. 9 in New material of Incadelphys antiquus (Pucadelphyda, Metatheria, Mammalia) from the early Palaeocene of Bolivia reveals phylogenetic affinities with enigmatic North and South American metatherians

FIG. 9. — Incadelphys antiquus, MHNC 13906: B, right lateral view of the anterior skull. Scale bar: 5 mm.

FIG. 9 in New material of Incadelphys antiquus (Pucadelphyda, Metatheria, Mammalia) from the early Palaeocene of Bolivia reveals phylogenetic affinities with enigmatic North and South American metatherians

FIG. 9. — Incadelphys antiquus, MHNC 13906: A, left lateral view of the anterior skull. Scale bar: 5 mm.

FIG. 8 in New material of Incadelphys antiquus (Pucadelphyda, Metatheria, Mammalia) from the early Palaeocene of Bolivia reveals phylogenetic affinities with enigmatic North and South American metatherians

FIG. 8. — Incadelphys antiquus, MHNC 13906: A, dorsal view of the anterior skull; B, ventral view of the anterior skull. Scale bar: 5 mm.

FIG. 6 in New material of Incadelphys antiquus (Pucadelphyda, Metatheria, Mammalia) from the early Palaeocene of Bolivia reveals phylogenetic affinities with enigmatic North and South American metatherians

FIG. 6. — Incadelphys antiquus, MHNC 13906: A, stereopair of the left lower tooth row in occlusal view; B, stereopair of the right lower tooth row in occlusal view. Scale bar: 5 mm.

FIG. 4 in New material of Incadelphys antiquus (Pucadelphyda, Metatheria, Mammalia) from the early Palaeocene of Bolivia reveals phylogenetic affinities with enigmatic North and South American metatherians

FIG. 4. — Incadelphys antiquus, MHNC 13906: B, stereopair of the left maxilla and upper tooth row in occlusal view. Scale bar: 5 mm.

FIG. 5 in New material of Incadelphys antiquus (Pucadelphyda, Metatheria, Mammalia) from the early Palaeocene of Bolivia reveals phylogenetic affinities with enigmatic North and South American metatherians

FIG. 5. — Incadelphys antiquus, MHNC 13906:A, lateral view of the right upper tooth row; B, lateral view of the left upper tooth row. Scale bar: 5 mm.

FIG. 4 in New material of Incadelphys antiquus (Pucadelphyda, Metatheria, Mammalia) from the early Palaeocene of Bolivia reveals phylogenetic affinities with enigmatic North and South American metatherians

FIG. 4. — Incadelphys antiquus, MHNC 13906: A, stereopair of the right maxilla and upper tooth row in occlusal view. Scale bar: 5 mm.