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FIG. 16 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 16. Dromiciops gliroides ZIUT HL 19 mm (Microbiotheriidae, Microbiotheria), juvenile specimen, stained coronal sections in rostrocaudal order. Effects of shrinkage during histological processing obvious (e.g., size of maxillary nerve relative to its sulcus) A, Presphenoid-basisphenoid synchondrosis, large pterygopalatine fissure provides passage for nerve of pterygoid canal (asterisk) (s. 25.02.02). B, Veins of rostral branches of transverse canals form confluent, with canals largely incorporated into rostral portion of transverse basicranial sinus. Cells within sinus nondiagnosable but probably erythroid (s. 28.03.01; cf. fig. 33A). C, Rostral branches of transverse canals present and large (s. 29.01.03). D, Entrance to carotid canal (double-headed arrow) (s. 30.04.01). E, Anastomosis of internal carotid vein with basicranial venous plexus (s. 31.01.01). F, Same anastomosis in closeup, typical plexiform arrangement (s. 31.03.02). Key: AS, alisphenoid; astp, tympanic process of alisphenoid; at, auditory tube; BS, basisphenoid; bvp, basicranial venous plexus; cc, carotid canal; cs, cavernous sinus; EC, ectotympanic; ejv, external jugular vein; encg, endocranial carotid groove; hp, hypophysis; ica, internal carotid artery; icv, internal carotid vein; junc + con, junction containing confluence; lpn, lesser petrosal nerve; mxn, maxillary nerve; nph, nasopharynx; oa, ophthalmic artery; on, ophthalmic nerve;
FIG. 14 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 14. Caenolestes sp. IANIGLA uncatalogued (Caenolestidae, Paucituberculata), adult caudal cranium, selected coronal segments in rostrocaudal order (data source, table 2). In A and B, rostral branches of transverse canals and rostral portion of transverse basisphenoid sinus raise separate eminences on dorsal surface of bony endocranial floor. Large aperture (single asterisk), possibly related to a vessel communicating with ophthalmic veins, opens into rostral portion of sinus. In C and D, hypophyseal canaliculi (feature 1) communicate with cavernous sinus (hypophyseal fossa). In F and G, double asterisks identify trackways for vessels apparently associated with ventral petrosal sinus. Key: AS, alisphenoid; astp, tympanic process of alisphenoid; bcf, basicapsular fenestra; BO, basioccipital; BS, basisphenoid; cbf, foramen of caudal branch of transverse canal; cbtc, caudal branch of transverse canal; cc, carotid canal; chf, condylohypoglossal foramen (nonspecific); co, cochlea; ctbs, caudal portion of transverse basisphenoid sinus; EC, ectotympanic; encf, endocranial
FIG. 26. Dasyurus hallucatus AMNH M-16033 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 26. Dasyurus hallucatus AMNH M-16033 (Dasyuridae, Dasyuromorphia), adult caudal cranium in ventral view. Note large size of piriform fenestra, acting as exit aperture for mandibular nerve in absence of separate foramen ovale. Exocranial aperture of carotid canal cannot be seen adequately in this perspective. Key: astp, tympanic process of alisphenoid; bjs, basijugular sulcus; BO, basioccipital bone; BS, basisphenoid; cchf, caudal condylohypoglossal foramen; eam, external acoustic meatus; evpf, foramen for extracranial continuation of ventral petrosal sinus; excf, exocranial carotid foramen; io + pf, incisura ovalis and piriform fenestra (undivided); jf, jugular foramen; pglf, postglenoid foramen; pr, promontorium of petrosal; ptpf, pterygopalatine fissure; rchf, rostral condylohypoglossal foramen; stmf, stylomastoid foramen; tcf, transverse canal foramen.
FIG. 13 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 13. Caenolestes sp. IANIGLA uncatalogued (Caenolestidae, Paucituberculata), 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 left lateral (rev.); D, endocranial floor, based on A (rev.); E, intact caudal cranium, reconstructed trackways of transverse canal branches superimposed on ventral surface. In D, note small apertures (hypophyseal canaliculi) on periphery of hypophyseal fossa in addition to caudal branch foramen. Inferred streamlines (small black arrows) suggest communication between caudal branch vein, internal carotid veins, and cavernous sinus. Large apertures opening into rostral portion of transverse basisphenoid sinus (asterisks) are of unknown function but may relate to track of nearby ophthalmic veins rather than craniopharyngeal canal (cf. Perameles, fig. 32C). Key: bcf, basicapsular fenestra; cbf, caudal branch foramen; cbtc/ le, caudal branch of transverse canal inside lateral extension of transverse basisphenoid sinus; cbv, caudal
FIG. 12. Monodelphis domestica NMB c. III.777 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 12. Monodelphis domestica NMB c. III.777 (Didelphidae, Didelphimorphia), adult caudal cranium, selected coronal segments in rostrocaudal order (data source, table 2). In A–E, note general resemblance to Caenolestes in mesocranial organization (figs. 13A; 14A, B), with prominent transverse canals and basisphenoid sinus, multiple openings in endocranial floor. Asterisk identifies either an internal compartment within transverse basisphenoid sinus, or an improbably large craniopharyngeal canal. In D–G, note partial tubes or grooves for caudal branches of transverse canal, absent in Didelphis (fig. 5), leading to caudal branch foramen (arrow). Key: AS, alisphenoid; astp, alisphenoid tympanic process; cbtc, caudal branch of transverse canal; cc, carotid canal; ctbs, caudal portion of transverse basisphenoid sinus; encf, endocranial carotid foramen; encg, endocranial carorid groove; excf, exocranial carotid foramen; hpf, hypophyseal fossa; junc, junction of transverse canals; le, lateral extension of transverse basisphenoid sinus; mxns, sulcus for maxillary nerve; onvs, sulcus for ophthalmic neurovascular array; pf, piriform fenestra; ptc, pterygoid canal; rbtc, rostral branch of transverse canal; rtbs, rostral portion of transverse basisphenoid sinus; SQ, squamosal; tcf, transverse canal foramen; tmc, tympanic cavity.
FIG. 6 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 6. In A, Philander opossum AMNH M-28329 (Didelphidae, Didelphimorphia), adult specimen, ventral view. In B, P. opossum AMNH M-266385, juvenile specimen, caudal cranium in oblique caudolateral view. Note relatively small size of transverse canal foramina in both specimens compared with exocranial carotid foramen (double foramina on right side in A, asterisk). Note also joint foramen for external continuation of ventral petrosal sinus and internal jugular vein. Key: astp, tympanic process of alisphenoid; bjs, basijugular sulcus; BO, basioccipital; BS, basisphenoid; cchf, caudal condylohypoglossal foramen; cspf, craniospinal
FIG. 33 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 33. Perameles sp. ZIUT HL 17.5 mm (Peramelidae, Peramelemorphia), perinatal specimen, stained coronal sections in rostrocaudal order. A, Rostral branch veins forming mesocranial confluence; active signs of hematopoiesis in compound structure with rostral portion of transverse basicranial sinus (ss. 39.03.02, 40.03.03). B–D, Trunk of transverse canal vein successively receiving its source vessels—caudal and rostral branch veins (s. 40.03.03, 42.03.04, 43.02.04). E, Notable size difference between basicranial venous plexus and ventral petrosal sinus (s. 51.01.04). F, Ventral petrosal sinus descending into its extracranial foramen (s. 59.01.02). G, Anastomosis of extracranial continuation of ventral petrosal sinus with internal jugular vein, ventral to jugular foramen (s. 62.02.02). H, Caudal branch of transverse canal vein joining internal carotid vein, and possible example of peritrigeminal emissarium (as defined by Aplin, 1990) running from foramen ovale to maxillary vein (single asterisk, s. 41.04.03). I, J, Condylar plexus including extracranial continuation of ventral petrosal sinus (double asterisks), internal vertebral venous plexus inside vertebral canal, sigmoid sinus/vertebral vein anastomosis, vertebral artery passing through foramen transversarium of atlas and travelling beneath brainstem (ss. 67.01.02, 70.02.02). In A, left side of confluent's lumen slightly enhanced in order to differentiate it from background. Cells within confluent include differentiating erythrocytes; those populating walls of bony canal are osteoclasts and osteoblasts. Other cell types in lumen are consistent with active hematopoiesis in highly cellular bone marrow (cf. Cline and Maronpot, 1985; Young et al., 2014). Key: AC, auditory capsule; ahp, adenohypophysis; AS, alisphenoid; at, auditory tube; bbs, basisphenoid-basioccipital synchondrosis; BO, basioccipital; C1, atlas vertebra; C2, axis vertebra; cbv, caudal branch of transverse vein; cdv, condylar emissary vein (not specific); cs, cavernous sinus; dcg, dorsal cervical ganglion; EO, exoccipital; evpf, foramen for extracranial continuation of ventral petrosal sinus; evps + ijv, extracranial continuation of ventral petrosal sinus joining internal jugular vein; GO, goniale; hgn, hypoglossal nerve; ica, internal carotid artery; icn, internal carotid nerve; icv, internal carotid vein; icv/bvp, internal carotid vein/ basicranial venous plexus; ivvp, internal vertebral venous plexus; jf, jugular foramen; lptm, lateral pterygoid muscle; MC, meckelian cartilage; MD, mandible; mdn, mandibular nerve; mptm, medial
FIG. 34. Notoryctes typhlops AMNH M–202103 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 34. Notoryctes typhlops AMNH M–202103 (Notoryctidae, Notoryctemorphia), adult caudal cranium in A, ventral, and B, oblique lateral views. In A, note internal subdivision within carotid groove (asterisk), assumed to separate internal carotid neurovascular bundle from channel for caudal branch of transverse canal. Key: astp, tympanic process of alisphenoid; BO, basioccipital; BS, basisphenoid; cspf, craniospinal foramen; eam, external acoustic meatus; evpf, extracranial extention of ventral petrosal sinus; evpf + jf, joint aperture for extracranial extension of ventral petrosal sinus and internal jugular vein; excf, exocranial carotid foramen; fm, foramen magnum; fo, foramen ovale; gf, mandibular fossa; jf, jugular foramen; ptpf, foramen for nerve of pterygoid canal; rchf + cchf, joint aperture for rostral and caudal condylohypoglossal foramina; sof, sphenooccipital fissure; stmf, stylomastoid foramen; tcf, transverse canal foramen; tmc, tympanic cavity.
FIG. 11. Monodelphis domestica ZIUT PND 12 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 11. Monodelphis domestica ZIUT PND 12, HL 8.5 (Didelphidae, Didelphimorphia), young postnatal specimen, stained coronal sections in rostrocaudal order. In A and B, note anastomotic link between internal carotid vein and presumptive trunk of transverse canal vein (ss. 22.03.03, 23.02.01). In C, note internal carotid vein/basicranial venous plexus lying medial to auditory capsule (s. 25.04.02); size diminished compared to appearance in A. In D, unnamed emissaria from ventral petrosal sinus (circled) passing through basicapsular
FIG. 9 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 9. Caluromys sp. ZIUT PND 77 (Didelphidae, Didelphimorphia), postnatal specimen, stained coronal sections in rostrocaudal order. A–C, sections through rostral portion of transverse basisphenoid sinus (ss. 71.02.02, 78.03.02, 79.03.02); D–F, sections through rostral end of tympanic cavity and carotid canal, showing rapid diminution of internal carotid vein (ss. 76.03.03, 77.02.03, 77.03.01). There is no evidence of trunk of transverse canal vein or its branches in expected positions. Internal carotid artery and vein still fill endocranial carotid grooves, even though shrinkage occurred during histological processing. Asterisk, branches of pharyngeal and pterygoid plexuses. Key: AC, auditory capsule; ahp, adenohypophysis; AS, alisphenoid; at, auditory tube; bbs, basisphenoid-basioccipital synchondrosis; BS, basisphenoid; bvp, basicranial venous plexus; cs, cavernous sinus; ctn, chorda tympani nerve; EC, ectotympanic; ejv, external jugular vein; encg, endocranial carotid groove; GO, goniale; ica, internal carotid artery; icn, internal carotid nerve; icv, internal carotid vein; MD, mandible; mdn, mandibular nerve; mxn, maxillary nerve; mxv, maxillary vein; oa, ophthalmic artery; on, ophthalmic nerve; ov, ophthalmic vein; pal, processus alaris; pglv, postglenoid vein; pvp, pterygoid venous plexus; spm, stylopharyngeus muscle; tg, trigeminal ganglion; tmc, tympanic cavity; tmm, tympanic membrane; ttm, tensor tympani muscle; vps, ventral petrosal sinus.
FIG. 3 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 3. Didelphis virginiana (Didelphidae, Didelphimorphia), representation of adult cephalic venous tree in ventral aspect, based on corrosion cast illustrations and descriptions by Dom et al. (1970: figs. 4, 5). Original authors' verified identifications (altered as necessary to conform to our terminology) are in regular type; italicized type and bold numbers represent additional identifications made for this paper (see key; color scheme as in fig. 1). Dorsal petrosal sinus, said to be present in this species (but see Shindo, 1915), is omitted. Craniopharyngeal canal, whatever its function, not depicted (foramen is present in Didelphis AMNH M-217731, fig. 2C). Right/left differences in vessel calibers probably represent natural variation or possibly distortions caused by latex injection. Dom et al. (1970: 494) state that "Plexus vertebralis internus ventralis appears to be the major pathway for drainage of the dural venous sinuses," but in our terminology this description should apply to vertebral vein per se, draining to craniospinal venous system. Internal plexus may instead be represented by relatively small vessels beneath brain stem. Key: con, confluence of transverse canal veins (rostral branches) (dark blue); cs, cavernous sinus (light gray); ejv, external jugular vein (dark gray); evps, extracranial continuation of ventral petrosal sinus (orange), here inferred to directly anastomose with vertebral vein; fv, facial vein (dark grey); icv + bvp, internal carotid vein and basicranial venous plexus (purple); ijv, internal jugular vein (light blue); mxv, maxillary vein (dark gray); ovp, ophthalmic venous plexus, including ophthalmic vein (dark gray); pglv, postglenoid (emissary) vein (dark grey); pvp, pterygoid venous plexus (aquamarine); tbs/le, rostral portion of transverse basicranial sinus and lateral extension (mottled green); tcv (rbv),
FIG. 30. Thylacinus cynocephalus NMB c.2526 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 30. Thylacinus cynocephalus NMB c.2526 (Thylacinidae, Dasyuromorphia), 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 vascular features; E, ventrolateral surface, intact caudal cranium; F, as in D, but with shape of hypophyseal fossa (yellow) separately depicted and resected to show position of endocarotid foramina. In A and B, carotid and transverse canals form complicated multiple union; see text and also figures 29 and 31. In C, no sulcus is in evidence to connect jugular foramen with sigmoid sinus trackway (which instead exclusively joins sulcus for vertebral vein), consistent with internal jugular vein reduced or absent. In D, single asterisks indicate small openings in transverse canals and other features, mostly representing connection points (interstitial canaliculi) with transverse basisphenoid sinus. In F, communication between caudal branch veins, cavernous sinus, and internal carotid vein is suggested but its actual soft-tissue form cannot be reconstructed. Because both CRVs originate from CS/ICV they are not considered to form a confluence like that of RBVs. Internal carotid artery would have passed rostrally out of endocranial carotid foramen as ophthalmic artery. Other features in D: a, large connector joining transverse canal to accessory ventral petrosal sinus; b, intercanal branch, significance unknown; c, notochord (or transclival canal); d, reduced left accessory ventral petrosal sinus, partly hidden by carotid canal (compare right side); e, canaliculus linking c and d. Key: avpsc, canal for right accessory ventral petrosal sinus; cbtc, caudal branch of transverse canal (pathway); cc, carotid canal; cchc, caudal condylohypoglossal
FIG. 2. Didelphis virginiana AMNH M-217731 in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 2. Didelphis virginiana AMNH M-217731 (Didelphidae, Didelphimorphia), caudal cranium in A, ventral; B, oblique caudal, and C, endocranial views. In C, only one foramen (for internal carotid neurovascular bundle) opens into endocranial carotid groove, because caudal branch of transverse canal vein and its foramen are not present. Small aperture opening in rostral wall of hypophyseal fossa is cranio-
FIG. 1. A in Transverse Canal Foramen And Pericarotid Venous Network In Metatheria And Other Mammals
FIG. 1. A. Cartoon of pericarotid venous network (PCVN) in a hypothetical marsupial in lateral aspect, with chief components of network in bold type (color scheme also in fig. 3). In addition to transverse basisphenoid sinus, pneumatic sinuses (not separately named) in presphenoid and basioccipital are also frequently present. Six major venous connections involving PCVN components occur in most investigated marsupials at some stage of development and are indicated by italicized letters: a, cs → vps (e.g., Osphranter, fig. 20C); b, cs → icv (e.g., Perameles, fig. 33D); c, icv → bvp (e.g., Monodelphis, fig. 11C); d, rbv → rbv (confluence of antimeres) (e.g., Dromiciops, fig. 16B, C); e, icv → ttcv via cbv (e.g., Caenolestes, fig. 13D); f, ttcv → pvp (e.g., Caluromys, fig. 9E). Smaller accessory connections (e.g., hypophysial and interstitial canaliculi), not separately illustrated, vary considerably among taxa or individuals. Asterisk, craniopharyngeal canal, frequently patent in adult but
Fig. 9 in Cryptic Speciation And Characteristics Of The Transition Bias Following An Example Of The Cytb Gene In Palearctic Mammals
Fig. 9. Variation of summarized tv/ts-index in micro- (1) and macromammals (2) depending on nucleotide substitution level. Thick lines illustrate exponential approximation.
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. 8 in Cryptic Speciation And Characteristics Of The Transition Bias Following An Example Of The Cytb Gene In Palearctic Mammals
Fig. 8. Variation of transition (upper lines) and transversion (lower lines) frequencies accordingly to substitution frequencies level. Think lines are empirical data for each subfamily/family, thick ones — polynomial approximations of averaged data.
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. 2 in Cryptic Speciation And Characteristics Of The Transition Bias Following An Example Of The Cytb Gene In Palearctic Mammals
Fig. 2. Average frequencies of nucleotide substitutions frequencies (sub) and its standard errors of the three taxonomical levels in micromammals (black) and macromammals (gray).
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