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FIG. 20 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 20. Selection of section drawings of Oldfieldthomasia cf. debilitata AMNH-VP 28600, from AMNH-VP archives; right side (ss. 8, 13, 17, 32, and ss. 19–22; approximately twice natural size). Drawings are presented with original pencil shading, but gray tone has been added to each section to improve overall legibility. Annotations are by G. G. Simpson; most are concerned with his efforts to locate possible or probable sutures ("s") bounding adventitious elements Xa and Xp. (Annotations have been repositioned to facilitate reading.) As Simpson himself concluded, most candidate "sutures" turned out to better interpreted as breaks. Asterisk
FIG. 19. Dipodomys merriami AMNH-M 182081 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 19. Dipodomys merriami AMNH-M 182081, caudal cranium in left lateral (rev.) (A) and dorsal (B) aspects; certain sutures emphasized to improve visibility. Names for middle-ear spaces and sutural bounds after Webster (1975). Insignificant rod (small black arrows in A, suprmeat sp squ in B) running across surface of multipartite middle ear (hypotympanum, epitympanum, antrum) is all there is of squamous portion of squamosal in this heteromyid. Rod ends freely over external acoustic meatus, and is nowhere fused to cranial sidewall. In A, black asterisk lies on equally insignificant "body" of squamosal. Apparent suture (large white arrows) crossing caudal bulge of immense middle ear in A and B marks track of an internal "mastoid" septum, seen through translucent walls. Roof of external acoustic meatus is formed by ectotympanic, whose
FIG. 18. Lepus californicus AMNH-M 177068 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 18. Lepus californicus AMNH-M 177068, left side (rev.) of caudal cranium in ventral (A) and oblique rostrolateral (B) aspects, illustrating extent of rarefaction (fenestration) and several osteological features related to intracranial "joint" (ICJ) between rostral and caudal parts of cranium. Both to scale in B. Only features of direct relevance are labeled; for a detailed cranial osteology of lagomorphs, see Wible (2007). The leporid ICJ, conceived by Bramble (1989) as a linked series of narrow gaps (sphenooccipital synchondrosis/ foramen ovale/piriform fenestra/midcranial hiatus), divides the skull into rostral and caudal moieties. In life these gaps would be filled with dense connective tissues, including sutural soft tissues and, within sphenooccipital synchondrosis only, cartilage. Hiatal plate (black asterisk), filling dorsal part of midcranial hiatus in B, is a process of the interparietal; it projects under processus squamosus (squ pr), perhaps acting as a strengthening member in absence of a caudally extensive squamous squamosal. Among cranial elements
FIG. 16 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 16. Protypotherium sp. MLP 12-2780, caudal cranium in left lateral (A), dorsal (B), and caudal (C) aspects. All to scale in C. Caudal cranium of Protypotherium strongly resembles that of close relative Cochilius (fig. 13). Both of these interatherioids possessed large epitympanic sinuses, although thecal prominence is relatively less than in pachyrukhines. Few sutures can be identified on the caudal cranium of this particular specimen, but there is no reason to think that it differed from other notoungulates in sutural content or element composition. Note broken areas above posttemporal canal. Large size of foramina for rami temporales is noteworthy (size varies within taxon; cf. Sinclair, 1909, pl. III, fig. 2).
FIG. 13 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 13. Cochilius volens AMNH-VP 29651, stereopairs of left side of caudal cranium (with keys) in ventral (A), oblique ventrolateral (B), and dorsal (C) aspects, on this and subsequent pages. Shading convention as in fig. 8. Key to A and B: 1, squamoso-alisphenoid suture (preotic area); 2a/2b, squamoso-ectotympanic suture (incl. glaserian fissure; question mark indicates boundary in meatal area uncertain); 3, putative ectotympano-entotympanic suture; 4, exoccipito-ectotympanic suture; 5, petrotympano-squamosal suture (question mark indicates uncertain boundary); 6, squamoso-exoccipital suture (on paroccipital process). Key to C: 1, squamoso-parietal suture; 2, inferred squamoso-interparietal suture; 3, speculative reconstruction of parieto-interparietal suture (externally obliterated, actual confirmation unknown); 4, supraoccipito-(inter)parietal suture; 5, squamoso-zygomatic suture. In A, petrosal (asterisk) slightly exposed in basicapsular fissure due to
FIG. 15 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 15. Cochilius volens AMNH-VP 29651, four virtual coronal sections, left side (rev.), arranged in rostral to caudal order (for approximate locations of sectioning planes, see fig. 14A). All to scale in D. A, section through rostralmost end of bulla. Note fracturing (br) on medial surface of squamosal, which mimics a true suture. Arrow, discontinuity on rostral lip of bulla thought to represent nearly obliterated ectotympano-entotympanic suture. B passes through external acoustic meatus (note incus in epitympanic sinus, out of position post mortem). Arrow identifies caudal end of same groove seen in A. Except for petroso-bullar suture, other discontinuities are fractures. Asterisk identifies the large longitudinal fracture, seen on dorsal aspect of skull and positioned
FIG. 12 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 12. Paedotherium sp. MLP 52-IX-28-29, seven virtual horizontal sections on this and subsequent pages, rostral to right. All to scale in A. For orientation, rostral portion of cranium is shown in A but cropped out in succeeding sections. In A, note continuity between transverse sulcus and posttemporal canal (white asterisks) and fracture (br, not a suture) passing through squamosal bar. Sections B–G are in dorsoventral sequence, beginning with a plane passing slightly beneath cranial roof and progressing through occipital condyles. Contributions of major elements to tentorium osseum, thecal alae, and walls of the transverse sinus are easily distinguished. Note extensive caudolateral exposure of petrosal material, unusual in a notoungulate.
FIG. 14 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 14. Cochilius volens AMNH-VP 29651, three virtual parasagittal sections, left side, arranged in lateral to medial order on this and subsequent pages; rostral to right. All sections to scale in C. Diagram A indicates planes of sectioning in this figure and figure 15. B, section through external acoustic meatus; note incus in epitympanic sinus, out of position post mortem (also seen in fig. 15B). Although remodeling has erased most suture lines, still partly represented is suture between ectotympanic's rostral crus and preotic portion of squamosal (squamoso-ectotympanic suture, large white asterisk). Suture between caudal crus and retrotympanic process is no longer identifiable. Upper leader to mand fos (mandibular fossa) points to damage comparable to that seen in a similar place in Oldfieldthomasia AMNH-VP 28600 (feature 3 in fig. 1A); it is not evidence of a suture. C, section through towered tip of cochlear promontory. Simpson (1936) thought the small septum (small white asterisk) on rear wall of bulla conducted internal carotid in Oldfieldthomasia; in Cochilius this feature is better interpreted as conduit for tympanic nerve. D, section through medial aspect of middle ear; the part of the bulla articulating with petrosal is probably of entotympanic origin (see text). Rostralmost end of septum bullae in this section is confluent with bullar wall; more caudally it is distinct and connects with
FIG. 11 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 11. Paedotherium sp. MLP 52-IX-28-29, four virtual parasagittal sections, left auditory region, rostral to right. All to scale in B. Sections are in lateromedial sequence, beginning with slice through external acoustic meatus and progressing through medialmost part of the tympanic cavity and bullar wall (see locator diagram, fig. 10B). In A, single white asterisk, sulcus for transverse sinus, grooving medial aspect of theca; double white
FIG. 17 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 17. Protypotherium sp. MLP 12-2780, virtual coronal sections through caudal cranium, apparent right side: rostral (A), middle (B), and caudal (C) portions of middle ear (A and C rev.). All to scale in C. For approximate locations of sectioning planes, see fig. 16B. Although this matrix-embedded specimen is in good condition overall except for damage to thecal walls, CT resolution is relatively poor and fine details cannot be discriminated. Note well developed medial paratympanic cavity in A and B. In B, arrow identifies approximate position of suture between squamosally derived theca and caudal crus of ectotympanic, not evident in this section but clearly present in others, confirming that squamosal contributes the most to epitympanic sinus enclosure. In C, caudal part of highly trabeculated epitympanic sinus extends deep into retrotympanic process (asterisk). Suture between supraoccipital and overplated interparietal is obvious.
FIG. 9. Paedotherium MLP unnumbered2 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 9. Paedotherium MLP unnumbered2, stereopair views of isolated left auditory region (with keys) in lateral (A) and medial (B) aspects. Both to scale in A. Shading convention as in figure 8. Specimen not in life position (see fig. 8B for proper orientation). This specimen, evidently adult, is relatively complete except for loss of thecal spine (single asterisk in A) and damage to exoccipital. Many features can be identified that are otherwise very difficult to see in intact skulls. Sutural divisions can be detected where expected in most areas; however, squamosal and ectotympanic are smoothly continuous (probable position of boundary indicated by question marks). Ectotympanic probably forms most or all of meatal roof; entotympanic participation has not
FIG. 8 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 8. Paedotherium sp. AMNH-VP 45914, stereopair views of caudal cranium (with keys) in ventral (A), right lateral (B), right caudal (C), and dorsal (D) aspects, on this and subsequent pages. All to scale in A. In key diagrams, squamosal (including external aspect of epitympanic theca) is white; other elements colored differing shades of gray. Question marks indicate indistinct or inferred borders between squamosal theca and some other element; hachure indicates major breakage. Delicate ventrocaudal extremity of paroccipital process is usually broken off in pachyrukhine skulls (see Sinclair, 1909: pl. X, text-fig. 4). Key to A: 1, petroso-exoccipital suture; 2, basioccipito-bullar suture; 3, exoccipito-bullar suture (on paroccipital process); asterisk lies in foramen sometimes identified as "foramen lacerum medium" or "carotid canal" in notoungulate studies (but see Gabbert, 2004). Note small, cylindrical tympanostyloids situated within hyoid recesses. Unidentified large basioccipital foramina (multiple on specimen's left side) lying close to the midline are likely venous ports. Key to B: asterisk in dorsal midcranial hiatus. Klinocrany is obvious in this aspect; compare less derived Cochilius (fig. 13B). Note within dorsal hiatus the slight exposure of petrosal (question mark) and tentorial processes of supraoccipital and parietal. Key to C: 1, supraoccipito-thecal suture; 2, petroso-exoccipital suture.
FIG. 10 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 10. Paedotherium sp. MLP 52-IX-28-29. A is virtual coronal section through external auditory canal, medial to right. B and C locate planes of sectioning for sections illustrated in figures 10–12 (exposed matrix colored gray). On skulls, ectotympanic material appears to contribute to dorsal wall of external acoustic meatus, although no suture between ectotympanic and squamosal theca can be identified in adult and actual situation therefore uncertain. Note incus (out of position) and prominent anular bridge. Asterisk (*) below bulla indicates fracture (not ectotympano-entotympanic suture).
FIG. 7 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 7. Oldfieldthomasia cf. debilitata AMNH-VP 28600, left hemicranium, virtual coronal section; for approximate location see figure 1B; lateral to right. In this section, the only suture found on preserved part of lateral wall of auditory region is the squamoso-ectotympanic suture; retroarticular process formed jointly by squamosal and ectotympanic, with no evidence of additional element Xa. Petrosal material contributes modestly to medial margin of aditus to epitympanic sinus and therefore to thecal wall. Matrix (see fig. 1) filling epitympanic sinus has been partially removed to facilitate labeling. Auditory ossicles—fragments of malleus and incus—are out of position in aditus to epitympanic sinus.
FIG. 8 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 8. (Continued.) Precise positions of sutures on caudal aspect of skull notably vary among Paedotherium specimens because overgrowths (alae) from surrounding elements may develop in such a way that they produce internal or self-sutures, thereby falsely suggesting presence of separate elements (e.g., "mastoideo-posttympanic suture" between theca and petrosal in Hegetotherium mirabile AMNH-VP 9223; Sinclair [1909: 71, text-fig. 14]). In this specimen, probable position of lines of fusion among elements indicated by question marks. Externally, aperture of posttemporal canal actually lies at junction of three bone territories (petrosal, squamosal, and exoccipital), but how the aperture is framed in the adult varies. Petrosal also produces alae, which may overgrow the posttemporal dehiscence in such a way that in some specimens (as here) the aperture appears to be wholly encased by petrosal material. In pachyrukhines the paroccipital process is a broad flange that incorporates not only exoccipital material but also ectotympanic and petrosal. Size of individual contributions also varies (cf. figs. 9B, 12F, G). In AMNH-VP 45914, both ectotympanic and petrosal appear on lateral side. As usual there is no sutural or other identifiable boundary between supraoccipital and exoccipital. Key to D: 1, supraoccipito-thecal suture; 2, parieto-thecal suture (along squamosal bar); asterisk,?lateral interparietal (see text).
FIG. 6 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 6. Oldfieldthomasia cf. debilitata AMNH-VP 28600, left hemicranium, virtual parasagittal sections (for approximate locations see fig. 1A); rostral to right. All to scale in C. A and B, sections through lateral cranial wall and external acoustic meatus: single white asterisk, definite suture between rostral crus of ectotympanic and alleged element Xa; double white asterisks, fracture zone (not a suture) passing through mandibular
FIG. 5 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 5. Oldfieldthomasia cf. debilitata AMNH-VP 28600, right hemicranium, parasagittal sections (after Simpson, 1936: fig. 5; original magnification, cited as natural size); rostral to right. This plate includes the only section illustrations of elements Xa and Xp in Simpson's paper (ss. 10, 15, 20, and 23; cf. comparable virtual parasagittal sections in fig. 6). Neither of these putative elements passes tests for recognizing independent ossifications; see text and appendix 1. Key (Simpson's nomina in parentheses, if different from usages preferred here; periods in original omitted): caf, facial canal (canalis facialis); cpg, retroarticular canal/foramen (postglenoid canal); crty, crista tympanica; cty, tympanic cavity; ets, epitympanic sinus; fcbr, cerebral fossa; fsmp, stylomastoid foramen (foramen stylomastoideum primitivum); fg, glaserian fissure (fissura glaseri); fpg, retroarticular canal/foramen (postglenoid foramen); fpn, aditus of epitympanic sinus (foramen pneumaticum); fsm, stylomastoid foramen; fv, venous foramen; fvpl, posterolateral cerebral venous foramen; mae, external
FIG. 4 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 4. Caudal cranium of adult Nesodon ovinus, as illustrated and interpreted by Roth (1903: pl. III, fig. 2; original magnification, cited as approximately one-third natural size); lateral aspect. The label "serrial" is simply floated over the external acoustic meatus, with no bounding sutures indicated, making it impossible to evaluate Roth's (1903) reason for discriminating it as such. Roth's key (terminology translated, updated, and annotated as in fig. 3): 3, occipital condyle; 4, paroccipital process; 5, retrotympanic process; 6a, "pars mastoidea" (= caudal section of pars canalicularis, exposed on margin of posttemporal canal); 7, crista meatus; 9, glaserian fissure; 11, retroarticular process; 12, mandibular fossa; 13, articular tubercle; 14, "incisura mastoidea" (= cleft between crista meatus and retrotympanic process); 21, external acoustic meatus.
FIG. 3 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 3. Skull of an adult Pachyrukhos typicus, as illustrated and interpreted by Roth (1903: pl. I, fig. 6; original magnification, cited as approximately natural size); oblique dorsolateral aspect. Roth's key (translated), with controversial names in quotation marks, preferred name following in parentheses: 1, supraoccipital; 4, paroccipital process; 5, retrotympanic process; 6, "pars mastoidea" (= epitympanic theca); 6a, crista lateralis; 16, auditory bulla; 19, "sutura mastoideo-supraoccipitalis" (= supraoccipito-thecal suture); 21, external acoustic meatus; 36, parietal; 37, squamosal; 38, squamoso-parietal suture; 41, "sutura squamoso-mastoidea (escamo-serrial)" (= actually parieto-thecal suture in this specimen, as the squamosal bar has broken away; see text).
FIG. 2 in The Serrialis Bone, Interparietals, "X" Elements, Entotympanics, And The Composition Of The Notoungulate Caudal Cranium
FIG. 2. Skull of a specimen of Toxodon, apparently juvenile (after Roth, 1903: text-fig. A; scale not provided), showing extent of supposed serrialis (gray shading) and other novel ossifications according to boundaries implied by Roth's determinations of supposed sutures. In fact, none of the elements so delimited qualifies as an independent ossification (see text). Abbreviations according to original caption: a, b, elementum serriale = prosquamosum; c, elementum posttympanicum = quadratum; mae, external acoustic meatus; pa, parietale; qui, elementum zygoma = quatrojugale; so, supraocipitale; sq, squamosum; ty, tympanicum; 1, sutura squamoso-parietalis; 2, sutura squamoso-serrialis; 3, sutura zygoma-serrialis; 4, [not provided in original]; 5, sutura mastoideo-posttympanica; 6, incisura mastoidea; 7, fissura glaseri. Although this has yet to be demonstrated for Toxodon, some of these alleged sutures could be the result of Roth's misinterpreting borders related to thecal alae—thin sheets of bone formed by the exoccipital, supraoccipital, and petrosal which encroach onto the squamosal thecae and sometimes form self-sutures, to varying extents among notoungulate taxa (cf. Paedotherium, fig. 12B–F).
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