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FIG. 21. Trigonostylops wortmani MLP 52-X-5-98, partial rostrum with C and P1 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 21. Trigonostylops wortmani MLP 52-X-5-98, partial rostrum with C and P1 (roots only) and P2-4, M1, and M2 (crowns bilaterally present) in A, ventral; B, rostral; C, oblique dorsal; and D, left lateral views. In A, appearance of rostral end of palate implies but does not prove that it naturally furcated into relatively large canine-bearing processes, separated by an intervening cleft (arrow). A separate premaxillary element cannot be detected within this cleft, but specimen is distorted and actual appearance in life remains uncertain. In B, nasal aperture is dorsally damaged on both sides, but on specimen's left side its margin or sill seems to be essentially intact, especially ventrally. External wall of canine alveolus is smooth, with no evidence of a complex sutural surface for lateral maxillopremaxillary suture. Trigonostylops possessed nasals of normal proportionate length that extended to end of rostrum (B, C). In C, nasals and maxillae are not in contact along their shared borders (arrow), but this seems to be a consequence of postmortem separation of nasomaxillary sutures, not presence of intervening process of premaxilla. In D, note presence of 3–4 infraorbital foramina in a nearly linear arrangement.
FIG. 15 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 15. Astrapotherium magnum, epitympanic and extratympanic sinuses and their adituses. A, MACN A 8580, virtual horizontal slice through left side of skull in ventral aspect, compared to photo of same skull in similar orientation; B, same skull and aspect, stereopair; C, MAPBAR 5322, extratympanic aditus in right retroarticular process, as seen from within external acoustic meatus. In A, white arrow points to small epitympanic sinus, situated lateral to position of (damaged) fenestra vestibuli (1); sinus slightly inflates tympanic roof near margin of
FIG. 14. Astrapotherium magnum MACN A 8580 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 14. Astrapotherium magnum MACN A 8580, transverse segments through skull in rostrocaudal order, all to same scale. In A, B, continuous chambers related to massive frontal sinus expansion serve to isolate cranial cavity from skull's sidewalls (suggested by black arrows); C, successive segments through apparent right side, illustrating relationship of frontal sinus (1) to extratympanic aditus (2) and other chambers in retroarticular process (3); D, segment through fenestra vestibuli (4) and aditus of epitympanic sinus, fully prepared on right side (white arrow) but still blocked by matrix on left; E, F, segments through rear of tympanic cavity and foramen magnum, illustrating positions of temporal sinus, posttemporal canal, sulcus for?vertebral artery, lambdoidal process of petrosal, and paracondylar process (cf. Tetramerorhinus, fig. 40H). In B, C, white asterisks mark a large longitudinal conduit within cranial sidewall that corresponds, in other SANUs, to trackway of temporal sinus. But this feature also communicates with spaces that open into retroarticular process' aditus, indicating that both kinds of sinuses (pneumatic and venous) were housed in same bony chambers (see text).
FIG. 13. Astrapotherium magnum MACN A 8580 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 13. Astrapotherium magnum MACN A 8580, parasagittal segments through left side of caudal cranium, sequence lateral to medial. In A, inset passes through skull just medial to the complete segment; note small epitympanic sinus and proximity of aditus of extratympanic sinus (see also fig. 27). In B, asterisk marks lambdoidal process of petrosal also seen in inset (see also fig. 14F). In C, D, curved black arrows indicate continuity between frontal sinus and air spaces throughout skull; note relatively small size of braincase. Cobbly appearance of bone and matrix in nasal region is due to artifacts in CT data. Premaxillae missing postmortem in this specimen (but see fig. 4).
FIG. 19 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 19. Differences between epitympanic and extratympanic sinuses and other features in three members of comparative set: Trigonostylops wortmani AMNH VP-28700, Cochilius volvens AMNH VP-29651, and Tetramerorhinus lucarius AMNH VP-9245. Top row: Selected segments through external acoustic meatus; note differences in scale. In each segment, dashed line indicates inferred position of tympanic membrane in life. Key (structures not exhaustively identified on each segment): 1, to aditus of extratympanic sinus (aperture not in plane of segment); 2, aditus of epitympanic sinus; 3, fenestra vestibuli; 4, ectotympanic; 5, cochlea; 6, tympanic membrane (inferred); 7,?sulcus for internal carotid artery on caudal portion of promontorium. Note that adituses of epitympanic sinuses of notoungulate Cochilius and litoptern Tetramerorhinus are located
FIG. 12 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 12. Trigonostylops wortmani AMNH VP-28700, transverse segments through skull (on this and following two pages) in rostrocaudal sequence and at different scales. In A–C, note artificial entrance (asterisk) into right palatal diverticulum (filled with matrix in B), aditus of palatal diverticulum opening into nasal cavity, and palatal alate process (left side broken). In D–G, note frontal sinus in sagittal crest, matrix-filled extratympanic sinus occupying lateral sidewall of skull, and preparation artefacts (asterisks) occupying original positions of tympanic cavity and rostral carotid foramen. In H–L, note artificially widened aditus of extratympanic sinus, reconstructed position of tympanic membrane (double-
FIG. 42 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 42. Visual guide for scoring certain matrix characters. "Slashed zero" symbol indicates "absent" character state. C113, Hypoglossal foramen (-ina): 0, located nearer to occipital condyle than to jugular area (e.g., Scalibrinitherium bravardi MACN Pv 13082); 1, hypoglossal foramen closer to jugular area than to condyle (e.g., Astraponotus sp. MPEF PV 1084); 2, hypoglossal foramen opens directly into jugular area (Trigonostylops wortmani AMNH VP-28700). C157 (new), Facial process of premaxilla: 0, present (e.g., Mesotherium angustirostrum MACN Pv 6040); 1, absent (e.g., Astrapotherium magnum AMNH VP-9278; Trigonostylops wortmani MLP 52-X-5-98) . C158 (new), Diverging canines (upper and lowers): 0, canines diverging at an angle less than 70° in front view (e.g., Eoastrapostylops riolorense PVL 4216; Phenacodus primaevus MACN Pv 18808, cast of YPM PU 14864); 1, divergence equal to or larger than 70° (e.g., Astrapotherium FMNH 13172; Trigonostylops MPEF PV 5483 [right canine reconstructed]). In the case of C159 (new), consult micrographs in original publication (Lindenau, 2005) for proper scoring.
FIG. 9 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 9. Meniscotherium chamense AMNH VP-4412, Cochilius volvens AMNH VP-29651, and Tetramerorhinus lucarius AMNH VP-9245, reconstructions of brain endocast and endocranial vasculature in dorsal, right lateral, ventral, and caudal views (on this and facing page). For neural features, see text. Key: 1, posttemporal sulcus/canal (for vasa diploetica magna); 2, temporal sulcus/canal (for temporal sinus); 3, parietosquamosal canals (for parietosquamosal vessels); 4, hypoglossal canal (for condylar emissary vein); 5, sulcus for vertebral vein; 6, sulcus for ventral petrosal sinus (may include part of cavernous sinus); 7, sulcus for sigmoid sinus (may include proximal internal jugular vein); 8, retroarticular sulcus/ canal (for retroarticular emissary vein); 9, cranioorbital sulcus/canal (for cranioorbital sinus); a, sulcus for transverse sinus or sinus communicans; b, accessory lacunae of transverse sinus (few in number, not reconstructed in detail); c, sulcus for dorsal sagittal sinus; d, ethmoidal canal (for ethmoidal neurovascular bundle); e,?dorsal petrosal sinus; f, petromastoid canaliculus (for parafloccular vein); g, transclival
FIG. 10. Ceratotherium simum AMNH M-51882, Tapirus indicus AMNH M-200300 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 10. Ceratotherium simum AMNH M-51882, Tapirus indicus AMNH M-200300, and Equus caballus AMNH M-204155, reconstructions of endocranial vasculature in dorsal, right lateral, ventral, and caudal views (on this and facing page). Specimens of C . simum and E . caballus are juveniles. For neural features, see text. As in previous figures in this series, vascular-related features are uniformly colored (gray), with exception of arteria diploetica magna (= caudal meningeal artery) (red) in Equus only. Posttemporal trackway contents in other taxa are grouped nonspecifically as vasa diploetica magna. See text. Key: 1, posttemporal sulcus/canal (for vasa diploetica magna); 2, temporal sulcus/canal (for temporal sinus); 3, parietosquamosal sulcus/canal (for parietosquamosal vessels); 4, hypoglossal canal (for condylar emissary vein); 5, sulcus for vertebral vein; 6, sulcus for ventral petrosal sinus (may include part of cavernous sinus); 7, sulcus for sigmoid sinus (may include proximal internal jugular vein); 8, retroarticular sulcus/canal (for retroarticular emissary vein); 9, cranioorbital sulcus/canal (for cranioorbital vessels); 10, sulcus for?ophthalmic vein; 11, sulcus for?caudal rhinencephalic vein; a, sulcus for transverse sinus or sinus communi-
FIG. 11 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 11. Trigonostylops wortmani AMNH VP-28700, parasagittal segments through left side of caudal cranium, sequence lateral to medial. In A, note preparation artifacts (gaps in matrix) in tympanic cavity and aditus of extratympanic sinus (asterisks); in B, distinct crista tympanica, track of retroarticular vein departing temporal sinus, and area of petrosal-squamosal fusion (arrows); in C, hypoglossal canal in rear wall of basicapsular fenestra, notch in ectotympanic for carotid artery, and cranial cavity exposed by preparation in endocranial wall of extratympanic sinus. Key: 1, aqueductus cochleae; 2, cavum supracochleare.
FIG. 8 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 8. Trigonostylops wortmani AMNH VP-28700 and Astrapotherium magnum MACN A 8580, reconstructions of brain endocast and endocranial vasculature in dorsal, right lateral, ventral, and caudal views (on this and facing page). Olfactory portion of brain mostly missing in Trigonostylops specimen; ventral petrosal sinus areas damaged in both. For neural features, see text. Top row: In vascular organization Trigonostylops resembles Meniscotherium and Tetramerorhinus (fig. 9) more than highly derived Astrapotherium. Despite unusual position in this taxon, hypoglossal canal communicates with sulcus for vertebral vein as in other SANUs. Small connector between transverse sinuses, possibly morphological equivalent of sinus communicans, was probably accompanied by other channels that left no impressions. Number and distribution of parietosquamosal canals is also noteworthy. Feature (single black asterisk in ventral view) seen on hindbrain/spinal cord is vascular, not neural, and composed of segmental vessels of uncertain homology. They drain from interior of occipital condyles toward endocast surface, and are therefore presumably tributary to vertebral venous system. Bottom row: Because of damage and scale of pneumatization, several expected channels, including
FIG. 7 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 7. Homalodotherium sp. MPM PV 17490, reconstructions of brain endocast and endocranial vasculature in A, dorsal; B, right lateral; C, ventral; and D, caudal views (on this and facing page). For neural features see text. Where possible, interpretations were checked against Patterson's (1937) latex endocast of Homalodotherium segoviae FMNH P13092. Morphological divisions of inferred dural sinuses and their connectors are color-coded to aid interpretation of similarly positioned (but uncolored) structures in figures 8-10. Key: yellow, dorsal sagittal sinus and transverse sinuses, including accessory lacunae of latter; brown, vasa diploetica magna (whether both arteria and vena diploetica magna were present is not determinable); green, temporal sinus and connectors (retroarticular vein, cranioorbital sinus, parietosquamosal veins); blue, sigmoid sinus and connectors (petrosal sinuses and condylar, vertebral, and internal jugular veins). In C, red indicates apparent trackways for putative rostral cerebral and vertebral arteries. Inset, segment at level of hypoglossal canals illustrating large sulci for inferred vertebral arteries (white asterisks). In this and following vascular reconstructions, CT resolution inadequate to detect separate trackways of arteries that typically accompany veins (e.g., arteria diploetica magna in posttemporal canal; meningeal arterial branches in parietosquamosal canals; cranioorbital artery accompanying cranioorbital sinus). Left side of specimen's basicranium is damaged, which accounts for incomplete reconstruction of some vessels (e.g., sigmoid sinus). Most vessels reconstructed from shallow sulci or other indicators and sizes cannot be considered dimensionally accurate. Supposed accessory vascular tissues (yellow) in extraneural space dorsal to cerebellum require further study (see text).
FIG. 2 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 2. Trigonostylops wortmani AMNH VP-28700, maxillary dentition (stereopairs): A, right side; B, left side. Defect on right side of palate is an artificial opening into palatal diverticulum (see text and fig. 12A).
FIG. 5 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 5. Equus caballus, principal vasculature of brain and endocranium (on this and facing page). After Leisering (1888: pl. 30, figs. 1, 3; pl. 31, figs. 1, 2). A, Arterial network of brain, ventral aspect; B, deep dissection of right basicranium and infratemporal fossa in oblique lateral aspect, showing preendocranial course of internal carotid artery and principal nerves. C, deep dissection of ventral aspect of head and rostral cervicals,
FIG. 4 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 4. Astrapotherium magnum AMNH VP-9278: A, ventral, B, dorsal, C, caudal, and D, right lateral views. Specimen is somewhat distorted and partly restored (see text). Apart from size, most noticeable external differences between Astrapotherium and Trigonostylops are concentrated in facial region (relative size of nasals and nasal cavity, rostral dentition, scale of pneumatization). Basicranial regions are also markedly different, but they share unusual derived features not found in non-astrapothere SANUs (see figs. 13–15, 27).
FIG. 1 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 1. Trigonostylops wortmani AMNH VP-28700, skull: A, ventral; B, dorsal; C, right lateral; D, left lateral; and E, caudal views (on this and facing page). In A, rectangle encloses palatal alate process (see fig. 20). For details of orbital and basicranial regions, see figures 25 and 26.
FIG. 3 in Cranial Morphology And Phylogenetic Relationships Of Trigonostylops Wortmani, An Eocene South American Native Ungulate
FIG. 3. Trigonostylops wortmani. Top: Mandible (MPEF PV 5483) with dentition: A, occlusal, B, rostral (mesial), and C, right lateral views. D, Reconstructed skull in right lateral aspect, chiefly based on AMNH 28700, MLP 52-X-5-98, and MPEF PV 5483.
Figure 7. The strict consensus tree obtained from the parsimony analysis with 35 in Descriptions and phylogenetic relationships of two new genera and four new species of Oligo-Miocene waterfowl (Aves: Anatidae) from Australia
Figure 7. The strict consensus tree obtained from the parsimony analysis with 35 characters ordered. Support values above lines at each node show bootstrap> 50% and Bayesian credibility values> 70% (100% = *). Values below lines are numbers of unambiguous synapomorphies for each node. Clades A, B, and C are referred to in text and Table 4.
Figure 5 in Descriptions and phylogenetic relationships of two new genera and four new species of Oligo-Miocene waterfowl (Aves: Anatidae) from Australia
Figure 5. Left humeri of tadornines in caudal (A, B), cranial (C, D) and ventral (E, F) views: A, C, E, modern T. tadornoides SAM B.39591; and B, D, F, Australotadorna alecwilsoni (SAM P.43141). The arrow points to the planar caudoventral margin of the bicipital crest compared to the angled margin in Tadorna. Scale bar = 10 mm. See main text for abbreviations.
Figure 2 in Descriptions and phylogenetic relationships of two new genera and four new species of Oligo-Miocene waterfowl (Aves: Anatidae) from Australia
Figure 2. Referred elements of Pinpanetta tedfordi: A–C, right coracoid SAM P.23477 in A, ventral; B, dorsal; and C, medial aspect; D, left tibiotarsus UCMP 56999 in anterior view; and E, left tarsometatarsus SAM P.24004 in dorsal aspect. Scale bars = 10 mm. See main text for abbreviations.
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