Otekaikea huata, Tanaka, Y & Fordyce, R, 2015

Tanaka, Y & Fordyce, R, 2015, A new Oligo-Miocene dolphin from New Zealand: Otekaikea huata expands diversity of the early Platanistoide, Palaeontologia Electronica (London, England) 9 (2), pp. 1-71 : 5-32

Otekaikea huata , new species

Figures 2-20 , Tables 1-3

zoobank.org/ CFD7D54E-00B2-40F1-830E-E60644595463

Diagnosis. Otekaikea huata is an odontocete with: a skull of medium size (bizygomatic width 265 mm, cranial length 222+ mm, likely condylobasal length 790+ mm); apical teeth procumbent with a large tusk (197+ mm in length, not including broken crown); more-posterior teeth near-homodont and polydont, with subconical crowns and at least one tooth with vestigial denticles; supraoccipital depressed; condyles prominent; zygomatic processes robust; and cervical vertebrae unfused. Otekaikea huata is slightly larger than the holotype of O. marplesi in terms of bizygomatic width (8 mm larger), maximum anteroposterior length of the periotic parallel to dorsal margin (7 mm larger) and also all measurements of preserved postcranial elements (each several mm larger). The one cheek-tooth known for O. marplesi has double roots, whereas the cheekteeth in O. huata are all single-rooted, including at least one with a vestigial longitudinal groove as if separating two fused roots. Otekaikea huata also differs from O. marplesi in: ascending process of the premaxilla is reduced, stopping at the level of the nasal (character 58); lateral expansion of the maxillae roofs the temporal fossa (character 101); nasal has an anterolateral process; fenestra rotunda of the periotic is very narrow and slit-like (character 174); caudal tympanic process of the periotic is posteroventrally projecting (character 178); and a prominent fold separates the infraspinous fossa and teres fossa on the scapula (character 236).

Etymology. From the Maori language: huata , a spear, alluding to the long tusk of the holotype specimen.

Material. Holotype only: OU 22306 ; the skull, right periotic and sigmoid process of left bulla, fused mandibles, 60 isolated conical-crowned teeth, atlas, axis, three other cervical vertebrae, 12 thoracic vertebrae, two lumbar vertebrae, eight ribs, scapulae, humeri, radii, an ulna, a carpal and nine digits. Collected by R.E. Fordyce, A. Grebneff, B. V.N. Black, and O. Sarll, 15 January and 12-13 February 1996; field number REF12-2-96-2.

Type locality. Informally-named Haughs’ Quarry , Hakataramea Valley , South Canterbury, New Zealand ( Figure1 ). See details above .

Horizon. A few centimeters below the diffuse Protula shellbed in the upper part of the Maerewhenua Member, Otekaike Limestone ( Figure 1 ). See details above.

TANAKA & FORDYCE: FOSSIL PLATANISTOID DOLPHIN

Age. Waitakian stage, Oligo/Miocene boundary, in the range 22.28 to 24.61 Ma. See details above.

General Description

Descriptions are based on the right or left side, whichever is more informative, with differences between them mentioned only if directional asymmetry is evident. Morphological terms follow Mead and Fordyce (2009) for the skull; postcranial terms mainly follow Flower (1885a), and Evans and de Lahunta (2013). Surface detail is lost in places from bioerosion, as noted when major. Some adherent concretionary carbonate remains.

Body size. The body size of Otekaikea huata can be inferred by the Pyenson and Sponberg (2011) formula for stem Platanistoidea , Log(L)=0.92* (log(BIZYG)-1.51)+2.49. BIZYG (bizygomatic width) of O. huata is 26.5 cm, giving a reconstructed body length of 2.57 m. As with O. marplesi , this is a similar body length to modern Platanista gangetica : 2.6 m in adult female, 2.2 m in adult male ( Jefferson et al., 2008). The gender of the type specimen of O. huata is uncertain. Closely related fossil species have comparable sizes: W. maerewhenua , bizygomatic width 24.4 cm (= length 2.39 m); O. marplesi , bizygomatic width 25.7 cm (= length around 2.5 m ( Tanaka and Fordyce, 2014). For O. huata , the minimum condylobasal length of 79.0+ cm would comprise around 30 % of the BIZYG-predicted body size of 2.57 m, implying a long rostrum and large cranium with a short trunk. Not enough postcranial material is known to provide independent evidence of body length.

Ontogenetic age. Skull sutures are mostly closed but distinct. All preserved vertebral epiphyses (in the atlas, axis, three cervical, 12 thoracic, and two lumbar vertebrae) are fused. These features suggest that O. huata is equivalent in ontogenetic age to stage VI, physically mature adult, in Stenella attenuata as proposed by Perrin (1975, p. 42). The skull lacks obvious roughened or eroded bone that might indicate a gerontic age. Otekaikea huata also has an ossified carpus, which supports an adult age class based on a flipper radiograph study on Phocoena sinus by Mellor et al. (2009).

Skull topography. The cranium is moderately complete dorsally ( Figures 2-9 ), while the separated rostrum is broken and represented by dorsal parts (porcelanous portions) of the isolated right and left premaxillae, the posterior end of the right maxilla, and anterior of the vomer with the attached posterior end of the left maxilla ( Figure 7 ). The separated rostral elements can be placed close to original articulation with the cranium, although minor burial-related distortion prevents exact matches that would allow full reassembly. The incomplete rostral left maxilla, which is sutured with the vomer and can be articulated with the cranium, preserves an elongate, flat, medially-dipping suture that matches a corresponding suture on the ventral surface of the premaxilla; a comparable suture is on the right. Both premaxillae (porcelanous parts) can be articulated with the flat sutures on the maxillae to indicate approximate life position on the rostrum (below). To reconstruct the skull, the right premaxilla was positioned to match the left; there was no independent evidence of bilateral asymmetry in the rostral sections of premaxilla. The face has a wide hexagonal shape in dorsal view, is flat anteriorly and rises steeply posteriorly. Compression slightly distorts the skull; additionally, the vertex is naturally skewed to left, as seen clearly in a posterior view, which shows the right nasal and frontal in the midline. Natural directional asymmetry affects the nasals, frontals, premaxillae, and maxillae. In lateral view ( Figures 6 and 8 ), the posterior part of the face has very steep medial and posterior walls involving both maxilla and premaxilla, and these walls form a strongly elevated but anteroposteriorly short vertex comprised of the maxillae, premaxillae, nasals, and frontals. The temporal fossa is mostly invisible in dorsal view because of the laterally expanded temporal crest. The orbit is long and weakly arched, with relatively thin antorbital and postorbital processes, above which the frontal is almost fully overlapped by the maxilla. The anteriorly broken external nares open from a subvertical

PALAEO- ELECTRONICA.ORG narial passage about level with the postorbital processes. The nuchal crest is not developed. The pterygoids and most of the palatines are missing.

Rostrum. Most of the anterior part of the rostrum, comprising the maxillae and ventral parts of the premaxillae, is not preserved. The long, straight dorsal remnants of premaxillae are reconstructed based on posterior connections with the maxillae (see below). This reconstruction is consistent with a narrow, parallel-sided and deep mesorostral groove, while the broken margins of the mandible suggest a narrow elongate Y-shaped lower jaw and, by implication, a narrow rostrum. The thick posterior part of the vomer ( Figure 7 ) sits medially, with sutured remnants of the maxillae indicating a posteriorly wide rostrum. Bone is damaged at the right antorbital notch, but the profiles of the lacrimojugal and adjacent frontal suggest an open

TANAKA & FORDYCE: FOSSIL PLATANISTOID DOLPHIN notch with a little-projecting lateral antorbital process.

The rostral porcelanous part of each premaxilla is incomplete posteriorly; this is interpreted as postmortem break when the cranium and rostrum separated before final burial. The elongate oblique suture on the maxilla is consistent with the premaxilla passing back to merge with the cranial parts of the premaxillary sac fossae that rise steeply to the vertex. Surficial structures include the rounded dorsal surface of the premaxilla, the remnant of a shallow premaxillary sulcus at the posterior end of the rostrum, and a steep-walled mesorostral groove (see Figure 8 ).

Premaxilla. The rostral and cranial parts of the premaxillae are present ( Figure 7 ), but separated post-mortem. Each bone is represented by the long, thin, posteriorly-narrowing, faintly bowed (barely concave laterally) porcelanous part of the rostral surface, each found separate from the cranium; more-ventral rostral parts of the premaxilla are lost. In life, the straight medial margin of each premaxilla would have partly roofed the mesorostral groove, converging toward each other anteriorly, as reconstructed in Figure 8 . The left premaxilla is distorted slightly outwards and down toward the rostral apex, probably a result of burial. Both ends are incomplete for each premaxilla, and bone is missing between the posterior of the rostral part and the premaxillary sac fossa on the cranium. Anteriorly, the rounded external surface on the rostral left premaxilla (511.5+ mm long) rises steeply, facing obliquely outwards. Posteriorly, the external surface becomes narrower and more abruptly rounded, and rotates gradually to face obliquely medially at the posterior apex of the porcelanous part. The posterior ~ 200 mm of the ventral surface of premaxilla is smooth at the suture for the underlying flat dorsal face of the maxilla; this flat suture on the maxilla is horizontal anteriorly on the rostrum but posteriorly it rotates to become steeply oblique by the level of the antorbital notch, facing medially and slightly dorsally. Here, each premaxilla narrows posteriorly, and is deflected slightly outwards. A broken narrow medial shelf on the left premaxilla, barely visible in dorsal view, is presumed to be part of the anteromedial sulcus, originally arising further posteriorly from the nowmissing premaxillary foramen.

Posteriorly, each premaxilla thickens, widens, and rises upward. In dorsal view, the lateral edge of the premaxillae is widest (125.1 mm) at the level of the nares. In spite of some compaction crushing, there is some directional asymmetry in the cranial parts of the premaxillae: 1, the right premaxilla is wider, lies further toward the midline, with a morelaterally flanged posterolateral plate than the left; 2, the right intra-premaxillary foramen, which opens dorsally between the maxilla and the posterolateral plate, is larger (elliptical; 10.6 mm long, 4.8 mm wide), and the lip of the foramen is formed mostly by the premaxilla, with maxilla contributing posteromedially. The left intra-premaxillary foramen opens almost the same level with the right, and it is smaller (also elliptical; 3.0 mm long, 1.2 mm wide; occluded by cemented matrix) and formed by the premaxilla; 3, the blunt right posteromedial splint reaches half way back along the nasal, bounded laterally by a slightly raised crest and vertical face of maxilla, whereas the left splint appears not to reach the nasofrontal suture and the adjacent maxilla is lower and more-nodular. The intra-premaxillary foramen is probably one of many foramina from the infraorbital complex, as discussed below.

Maxilla. The rostral part of the left maxilla is firmly sutured with the vomer (possibly cemented by matrix); the two bones can be placed close to their original positions against the cranium, with the maxilla close to the antorbital notch. The rostral part of the right maxilla, not attached to the vomer (but originally sutured), can be mirrored against the left. Dorsally, each maxilla has an elongate planar suture dipping obliquely ventromedially, formed by dense rather than cancellous bone, for the premaxillae ( Figures 7 and 8 ); the sutural face can be traced back onto the cranium. A small (2.6 mm diameter) dorsal infraorbital foramen opens bilaterally, facing outwards, toward the posterior of the rostral maxilla. Ventrally, the maxillae form a medially arched palate with flattened lateral margins at the base of the rostrum.

Posteriorly, the cranial part of the maxilla is overlaid by the premaxilla, with clear sutures with the frontal and lacrimal. In places (e.g., left near the position of the maxillary-premaxillary suture), the maxillary surface is dense and porcelanous; adjacent irregular cancellous surface is interpreted as (bio) eroded. About 25 mm medial to the right antorbital notch, the maxilla has an anteroposteriorly long, wide and smooth trough, presumably partly for the premaxilla, extending ~ 15 mm anterior and ~ 40 mm posterior to the level of the antorbital notch ( Figures 2.2 , 7.4 ). This trough has a steep lateral face that passes sharply at an elongate fold onto the flat supraorbital process of the maxilla.

On the braincase, dorsally, the cranial part of the maxilla covers most of the frontal except the lateral part of the postorbital process (right side; Figure 2 ). The maxilla on the supraorbital process is flat and smooth; it is not obviously thickened and it lacks a pneumatic maxillary crest. At the level of the right lacrimojugal, behind the antorbital notch, a small anterior dorsal infraorbital foramen (1.4 mm diameter) opens into a short oblique sulcus ( Figure 2.2 ). The region is damaged on the left. The posteromedial surface of the maxilla is very steep; at the level of the nasal/frontal border, two posterior dorsal infraorbital foramina open on each side, which are small and rounded (right side diameters: anterior, 3.4 mm; posterior, 4.6 mm), opening into sulci that run toward the posterolateral edge of the

TANAKA & FORDYCE: FOSSIL PLATANISTOID DOLPHIN maxilla. The squared apex of each maxilla meets the frontal medially and the supraoccipital posteriorly. Just lateral to the nasal and frontal, the medial border of the weakly curved crest on the right maxilla rises abruptly. At the anterior end of the nares, a small area of the maxilla is exposed medially; whether this was originally covered by now-lost premaxilla, or exposed to form a maxillary intrusion (sensu Arnold and Heinsohn, 1996) is uncertain.

In ventral view ( Figure 3 ), the better-preserved right maxilla forms the borders for two infraorbital foramina: a larger and more posteromedial foramen in the common position for odontocetes anterior to the antorbital ridge, and a smaller foramen a little anterolateral. The larger posterior foramen is bounded by frontal, palatine and maxilla, and opens anteriorly into a sulcus in the maxilla. The smaller anterior foramen is bounded mainly by maxilla and lacrimojugal, plus a small contribution of frontal; the lacrimojugal cleft is immediately lateral. It is likely that this smaller foramen was originally covered by maxilla that was lost postmortem, with the sulcus representing the infraorbital canal. On the type of Otekaikea marplesi , the single visible ventral infraorbital foramen is equivalent to the posterior foramen in O. huata .

Palatine. Ventrally, a vertical plate-like robust fragment of palatine underlies the maxilla ( Figure 7 ), slightly posteromedial to the infraorbital foramen, and connected with the frontal posteriorly. The palatine has a smooth lateral wall and, medially, a grooved vertical surface that contributes to the lateral wall of the nares.

Pterygoid. A fragment of pterygoid is lateral to the posterior end of the vomer, posterior to the nares. Otherwise the bone is lost. A facet for the medial lamina of the pterygoid covers the anterior two thirds of the basioccipital crest.

Nasal. The smooth, flat, asymmetrical nasals lie just behind, rather than roofing, the nares ( Figure 2 ). In anterior view, the nasals are dorsoventrally thick and nodular. The smaller left nasal is suboblong, transversely wide and anteroposteriorly narrow; the lateral and posterior margins are rounded, the anterior, straight. The sagittallyplaced larger right nasal is a rounded irregular triangle, transversely wide and anteroposteriorly longer than the left, and projected forward more than the left. Both have a prominently rounded medial corner and a semicircular posterior margin. The anterior edge of each nasal lies far posterior to the antorbital process, almost level with the posterior end of the postorbital process. The anteromedial angle projects strongly, while the posteromedial angle is displaced by the narial process of the frontal. Each nasal has a plate-like anterolateral process of the nasal [new term] laterally, ~ 20 mm long and ~ 4 mm wide, projecting forward to meet the posteromedial margin of the premaxilla, and separated from the main part of the nasal (anterior view) by a deep groove; the anterior margins of the two nasals form a W-shape. A slightly similar condition of the groove on the anterior surface of the nasal is seen on some kentriodontids ( Pithanodelphis cornutus Abel, 1905 , Delphinodon dividum True, 1912 and Kentriodon pernix Kellogg, 1927 ), but the grooves are much shallower and the anterolateral process less marked than in Otekaikea huata . An undescribed platanistoid OU 22126, from the Otekaike Limestone of Waitaki district (Fordyce, 2003), shows an anteroposteriorly thin nasal without a groove on the anterior surface, but the plate-like anterolateral process of the nasal projects well anteriorly. The internarial, nasofrontal, nasopremaxillary, and nasomaxillary sutures are deep and wide. The internarial suture is invaded by narrow fingers of frontal, forming the narial process. In

TANAKA & FORDYCE: FOSSIL PLATANISTOID DOLPHIN anterior view, the nasal is medially very thick but laterally thin, resulting in a triangular profile.

Ethmoid. The ethmoid is used in the sense of Mead and Fordyce (2009), but note that Ichishima (2011) suggested that the mesethmoid might be absent in odontocetes. Morphological terms of the ethmoid follow Godfrey (2013).

The profile of the ethmoid is uncertain but in the mesethmoid ridge (area for attachment of the mesorostral cartilage) of the vomer, there is a spongy bone with a foramen at the center ( Figure 4 ). The large void dorsal to the foramen may be for the missing or undeveloped area of the cribriform plate of the mesethmoid. On the posterodorsal part, the cribriform plate has a pair of large and strongly curved crescentic foramina. Between the foramina, there is a tubercle, which is assumed to be the mesethmoid.

Vomer. The robust rostral part of the vomer is broken and isolated from the cranium ( Figure 7 ). The mesorostral groove is anteriorly shallow and Ushaped, and posteriorly deep and more V-shaped. Its lateral wall is unexpectedly thick, thinning ventrally; the greatest bilateral width is 75.4 mm, with one side attaining a thickness of 27.6 mm, and the depth is around 30 mm. From the widest point, the vomer narrows rapidly, both anteriorly and posteriorly. Both ventrally and laterally, the vomer is covered by the maxilla cemented by matrix; the sutures are not fused. In ventral view ( Figures 7 and 8 ), the vomer carries two flat areas, which might be sutures for the maxilla (this feature occurs in an unfused young individual of Cephalorhynchus hectori and in Otekaikea marplesi ). Vomer is present below the olfactory region, as shown in broken section in anterior view. Posteriorly, the vomer covers the basisphenoid.

Lacrimojugal. The right lacrimal and jugal are fused without evident suture. There is no obvious lacrimal foramen or groove. A transversely long (36.3+ mm), thin (6.8 mm), and bar-like anterior margin of lacrimojugal contributes to the right antorbital notch ( Figure 3.2 ), and is separated from the rest of the bone by a lacrimojugal cleft. There is no evident base for the styloid process of the jugal. Medially, the anterior margin enlarges to become subspherical, sutured with the maxilla in a fossa (seen ventrally on the left maxilla) close to a prominent anterior infraorbital foramen. The lacrimojugal does not, however, reach the larger more-posterior infraorbital foramen. More posteriorly, the lacrimojugal is plate-like, as also implied by the slightly ridged planar suture for the missing lacrimojugal under the left antorbital process of the frontal.

Frontal. Each frontal contributes a large surface to the orbit and a small exposure at the vertex ( Figure 2 ). At the vertex, the frontals are bounded by the nasals, maxillae, and supraoccipital; there is a clear but irregular median interfrontal suture, not to be confused with a mollusc shell imprint (limpet homing mark?) on the right nasal. Each frontal is trapezoidal and has a long finger-like medial projection which forms a narial process. The frontals are asymmetrical: wider on the left and longer on the right. Each frontal contacts the parietal laterally. On the face, the left ascending process of the maxilla is lost and shows the profile of the postorbital process; the right process, which is almost covered by maxilla, appears not to project as far as the left,

PALAEO- ELECTRONICA.ORG judging from the profile of the bar-like jugal (above).

Ventrally, the frontal forms the anterodorsal wall of the braincase and most of the shallow and long orbit ( Figure 3 ). A weak preorbital ridge ( Fordyce, 1994) is indistinct at the antorbital process laterally, but becomes elevated and narrow toward the posterior margin of the narial passage. The frontal contributes the posterior and lateral margins of the infraorbital foramen. Medially, an oval presumed optic foramen (3.0 mm diameter) opens anterolaterally. The strongly posteriorly-concave postorbital ridge is sharp, high, and meets the postorbital process, which is a blunt, ventrally-projecting triangular shape as seen in lateral view. Between these ridges, the triangular roof of the orbit is anteroposteriorly long and dorsoventrally shallow. Foramina for the diploic vein are absent.

Orbitosphenoid. The orbitosphenoid ( Figure 3 ) is identified by the presumed ethmoid foramen, which opens 61-62 mm medial to the orbital margin, lateral and anterior relative to the region of the optic foramen. The foramen is surrounded by a smooth bone surface, presumed orbitosphenoid, in the roof of the orbit toward the optic foramen. No sutures are evident.

Parietal. The parietal forms the lateral wall of the braincase ( Figure 6 ) at the temporal fossa, which is strongly excavated medially and anteroposteriorly shorter than in Archaeoceti and stem Odontoceti , which have dorsally more open temporal fossa. Anteriorly, the parietal has a roughly rectangular thin supratemporal flange [new term], directed obliquely forward to encroach the frontal in the roof of the right temporal fossa. (The flange is lost on the left, but an exposed suture is apparent.) Such a flange is also clear in the undescribed specimen OU 22540 and is uncertainly present (the bone surface is damaged) in Otekaikea marplesi . Within the temporal fossa, the parietosquamosal suture is not clear in O. huata , and there is no obvious postparietal foramen. Parietal forms a ventral exposure between the basioccipital and squamosal, separating the foramen ovale from the cranial hiatus as in Waipatia maerewhenua and O. marplesi . The parietal-alisphenoid suture is clear on the left, between the foramen ovale and the base of the falciform process. "Foramen 1" sensu Fordyce (1994) is seen in O. huata further suggesting the position of parietal (see ventral details of the squamosal, TABLE 2. Measurements in mm of holotype, OU 22306, Otekaikea huata : periotic and tympanic bulla. Dimensions follow Fordyce et al. (2002), Perrin (1975), and Kasuya (1973). Measurements are rounded to the nearest 0.5 mm.

Periotic maximum anteroposterior length, from anterior apex of anterior process to apex of posterior process 44.0

maximum anteroposterior length parallel to dorsal margin 43.0

maximum dorsoventral depth anterior process, perpendicular to axis of periotic 15.0

length of anterior process, from anterior apex to level of posterior of mallear fossa 22.5

length of anterior process, from anterior apex of anterior process to level of anterior of pars cochlearis 14.5 in notch immediately lateral to fine ridge dorsoventral depth at fovea epitubaria length facet on posterior process point to point facial canal anteroposterior diameter maximum width of anterior process at base approximate anteroposterior length of pars cochlearis 1.5

17.5

2.0

11.5

18.0

approximate transverse width of pars cochlearis, from internal edge to fenestra ovalis 10.5 transverse width of periotic, internal face of pars cochlearis to apex of lateral tuberosity 22.0

length of posterior process of periotic 18.5

length of posterior process parallel to posterior profile/ steeply acute to long axis of body 18.0

Tympanic bulla width of sigmoid process height of sigmoid process below). The clear parietal-alisphenoid suture has foramen 1 on its junction with the squamosal, but otherwise the relationships of the foramen and sutures are obscure. The parietal contacts "foramen 2" sensu Fordyce (1994) posteriorly and the cranial hiatus medially.

Squamosal. Description is based mainly on the right. In dorsal view ( Figure 2 ), a wide (34.2 mm at the subtemporal crest) and long zygomatic process (90.1 mm) projects slightly laterally, posterior to the orbit. The anterior end is blunt and narrower than the posterior, and reaches to the level of the posterior end of the nasals, still distant from the postorbital process (14.6 mm gap between the processes, possibly widened by postmortem distortion). The temporal fossa is exposed widely in lateral view, is not obvious in dorsal view, and is widely open to posterior view. The temporal fossa has an unclear squamosal-parietal suture on its lateral wall on the right ( Figure 6 ). In lateral view, the anterior end of the squamosal is squared. On the lateral surface, a triangular, depressed neck muscle fossa (34.0 mm long) occupies midpoint of the zygomatic. This depression has been termed sternomastoid fossa in recent literature on fossil Cetacea , but the muscle insertions in extant species are

14.0

8.5

quite variable ( Schulte, 1916; Howell, 1927, 1930; Cotten et al., 2008), and the more-general term neck muscle fossa ( Fordyce, 1981, p. 1035) makes fewer assumptions about homologies.

Ventrally ( Figures 3 and 9 ), the zygomatic process has a wide and shallow mandibular fossa without an obvious anterior border, a slightly swollen lateral portion, and a strongly curved internal face. The postglenoid process is not thickened anteroposteriorly, nor markedly expanded laterally, and slopes obliquely posteroventrally. The large, prominent tympanosquamosal recess separates the zygomatic process from the falciform process, occupying the squamosal from the subtemporal crest to the anterior meatal crest and spiny process. Posteriorly, the recess descends down the medial margin of the postglenoid process, while posteromedially the recess deepens toward the spiny process. The periotic apposes the falciform process dorsally and anteriorly, separated by a small gap. In the articulated periotic, the position of the lateral tuberosity lies more ventrally than the margin of the falciform process ( Figure 11 ). The apex of the spiny process points forward toward the lateral tuberosity of the periotic, to bridge (pass ventrally to) the path of the middle sinus. An irregu- TABLE 3. Measurements in mm of holotype, OU 22306, Otekaikea huata : postcranial elements. Dimensions follow Fordyce et al. (2002) and Perrin (1975). Measurements are rounded to the nearest 0.5 mm. + means measurements of preserved distances of broken part.

Atlas maximum preserved length maximum preserved height maximum preserved width length of body height of anterior articular surface width of anterior articular surface height of posterior articular surface width of posterior articular surface

Axis maximum preserved height maximum preserved width length of body length of dens height of anterior articular surface width of anterior articular surface height of posterior articular surface width of posterior articular surface

4th cervical vertebra maximum preserved length maximum preserved height maximum preserved width length of neural spine height of neural spine length of body height of anterior articular surface width of anterior articular surface height of posterior articular surface width of posterior articular surface

5th cervical vertebra maximum preserved length maximum preserved height maximum preserved width length of body height of anterior articular surface width of anterior articular surface height of posterior articular surface width of posterior articular surface

6th cervical vertebra maximum preserved height maximum preserved width

63.0

70.5+

101.0+

60.0

48.5

90.0

47.5

87.5

76.0

129.0

55.5

23.0

36.0

88.5

49.0

54.5

39.5

88.0+

80.5+

79.0+

11.0+

28.0

47.0

45.5

40.5

46.5

28.5+

71.5+

98.0+

27.0

51.5

51.5

49.0

51.0

74.0+

85.5+

length of body height of anterior articular surface width of anterior articular surface height of posterior articular surface width of posterior articular surface

Thoracic vertebra 1 in Fig. 14

maximum preserved height maximum preserved width length of body

Thoracic vertebra 2 in Fig. 14

maximum preserved length maximum preserved height maximum preserved width length of neural spine height of neural spine length of body

Thoracic vertebra 3 in Fig. 14

maximum preserved length maximum preserved height maximum preserved width length of neural spine height of neural spine length of body

Thoracic vertebra 4 and 5 in Figure 14

maximum preserved length maximum preserved height maximum preserved width length of neural spine height of neural spine length of body

Thoracic vertebra 6 in Figure 14

maximum preserved height maximum preserved width length of body

Scapula maximum preserved length maximum preserved height length of acromion process length of glenoid fossa width of glenoid fossa depth of glenoid fossa

26.5

44.0+

47.0

47.0+

46.5+

66.0+

107.0+

32.0

49.0+

120.0+

89.0+

23.0+

49.5+

44.0

83.5+

128.0+

83.5+

56.5+

49.0+

58.0

73.5+

95.5+

81.5+

52.0+

16.0+

63.0

60.5+

244.0+

74.5

left right

284.0+ 328.0

118.0+ 224.0

79.0+ 95.5+

52.5 50.5

47.5 48.5

10.0 9.0

Humerus

Length narrowest length of shaft width of shaft at the narrowest point

Ulna

Length length without olecranon process narrowest length of shaft width of shaft at the narrowest point

Radius

Length narrowest length of shaft width of shaft at the narrowest point

Sternum preserved length preserved widest width preserved narrowest width lar transverse depression lies lateral to the spiny process, immediately in front of the anterior meatal crest and behind a small anterior transverse ridge sensu Fordyce (2002, p. 203); this may include the sigmoid fossa sensu Geisler et al. (2005, p. 16). The long base of the well-developed falciform process arises near the squamosal-alisphenoid suture at the subtemporal crest, and extends to the spiny process. There is no evidence that a lateral lamina was directed forward from the falciform process to bound the pterygoid sinus fossa. The falciform process is skewed medially to underlie the alisphenoid; its ventral tip has a small oblique facet probably for the outer lip of the bulla, while further dorsally the concave posterior face closely parallels the anterior process of the periotic. Laterally, the margin of the falciform process closely matches the anterior process and lateral tuberosity of the periotic; the spiny process fills the hiatus epitympanicus but with the periotic articulated it is apparent that a few mm of the spiny process is missing (there is incomplete contact posteriorly at the hiatus epitympanicus). The walls of the external auditory meatus form a narrow triangle medially, widening a little laterally, where the meatus undercuts the postglenoid process and passes into a groove that rises forward onto the lateral face of the squamosal ventral to the neck muscle fossa.

left right

153.0 151.0

55.0 56.0

39.5 41.5

left right

- 146.0

- 103.0

- 40.0

- 21.0

left right

115.0 120.0

34.0 37.0

26.5 27.5

97.0+

86.5+

58.0+

Associated with the latter groove is a small notch at the lateral end of the meatus. The anterior meatal crest is damaged; the posterior crest is arched transversely, with a facet on the posterior face at the suture for the posterior process of the bulla.

The anteroposteriorly long periotic fossa is hidden when the periotic is in situ; it includes anterior and posterior portions that are transversely wide and divided by a strong ridge. In the middle of the anterior portion, the foramen spinosum has two openings, anterior and posterior, which are separated by an elliptical tubercle, as also seen in Otekaikea marplesi . The anterior foramen spinosum is associated with a deep groove (the parietalsquamosal and parietal-alisphenoid sutures), which runs to the foramen ovale. The posterior portion of the periotic fossa is transversely wide (15.9 mm) and deep, and contains the suprameatal pit. When in situ, the periotic sits firmly in a single position in the periotic fossa, with the apex of the posterior process in close contact with the squamosal, medial to the post-tympanic process, but separated a few mm from the exoccipital. A conical space, which opens medially between the periotic and exoccipital, could be for part of the peribullary sinus. The squamosal is fissured, with several foramina, directly above the posterior process of the periotic. The ventrolateral apex of the post-tympanic process lacks obvious grooved sutures for the tip of the posterior process of the tympanic bulla, in contrast to the structure in Waipatia .

Basioccipital. Ventrally ( Figure 3 ), the basioccipital is a trapezoid. Anteriorly, the basisphenoid-basioccipital synchondrosis is fused. Each basioccipital crest gradually widens posteriorly, forms a gently curved ventral profile, and has an indistinct muscular tubercle on the medial surface of the crest. Between the crests, the basioccipital basin is trapezoidal and flat. The lateral border of the basioccipital, which is slightly excavated behind the carotid foramen, contributes to the anterior and medial margins of the cranial hiatus. It is not clear whether the exoccipital contributes to the posterodorsal part of the basioccipital crest medial to the jugular notch.

Supraoccipital and exoccipital. Posteriorly, the supraoccipital is rugose and roughly squared, and constricted by the temporal fossa. The nuchal crest is weakly developed. The dorsal condyloid fossa is deep and wide (60.9 mm) and curves over the condyle. Each occipital condyle has a smooth and small surface, is semicircular in lateral view, is less obviously curved horizontally (dorsal view), is somewhat elliptical in posterior view, and has a very short but distinct pedicle that is slightly narrower than the condyle. The right pedicle has a small foramen (2.7 mm diameter), on the ventral surface. The foramen magnum is a dorsally wider trapezoid, widely open, wider than high (37.9 mm wide and 29.0 mm height), and slightly arched dorsally; the profile may result from dorsoventral crushing. The intercondyloid notch is U-shaped. Laterally, the exoccipital is a thin rectangular plate. There is no distinct posterior sinus fossa here.

In ventral view, the exoccipital extends laterally almost as far as the adjacent post-tympanic process of the squamosal. The ventral surface is a smooth, transversely elongate, narrow, tabular to slightly depressed, tongue-shaped paroccipital process for stylohyal articulation. The rounded tip of the process extends medially to within about 5 mm of the basioccipital crest, and partly underlies the oblique narrow jugular notch (2.9 mm wide, 10.1 mm deep) that opens posterolaterally above the paroccipital process. The small hypoglossal foramen opens in the medial end of the jugular notch.

Alisphenoid. The large alisphenoid forms the posteromedial margin of the subtemporal crest. On the basicranium ( Figures 3 and 9 ), the alisphenoid is posterior to the frontal, anterior to the squamosal and basisphenoid, and lateral to the pharyngeal crest; there are clear sutures with the squamosal and parietal, but the contact with the frontal is broken. A blunt projection of alisphenoid extends back to the base of the falciform process, but does not form a posterolaterally-prolonged wedge. There is no distinct suture with the basisphenoid medially; the latter is indicated by the carotid foramen, but its lateral extent beyond the foramen is uncertain.

Anteriorly, the pterygoid sinus fossa comprises 3 small shallow depressions between the subtemporal crest, the large medially-placed foramen for the optic nerve, foramen rotundum and orbital fissure, and the foramen ovale. The foramen for the optic infundibulum is anteroposteriorly long and elliptical (31.4 mm long and 25.6 mm wide); it is bordered by alisphenoid posteriorly, basisphenoid medially, and frontal (and orbitosphenoid?) anteriorly. Just slightly posterior to the pterygoid sinus, a weak transverse ridge on the alisphenoid marks the groove for the mandibular nerve and associated vessels between the small foramen ovale (8.8 mm wide and 7.6 mm long) and the anterior end of the falciform process. The foramen ovale is complete, formed entirely by alisphenoid, which is apparently sutured with itself posteriorly. The alisphenoid is separated from the cranial hiatus by 25-26 mm of parietal and possibly by basisphenoid. A prominent rod-like projection of alisphenoid underlies the anterior border of the foramen ovale, prolonged toward the basisphenoid.

Basisphenoid. Anteriorly, the basisphenoidalvomer suture has two foramina, which open posteriorly into anteroposteriorly long shallow grooves ( Figure 3 ). The basisphenoid-basioccipital synchondrosis is fused. Laterally, a large and rounded carotid foramen (5.3 mm diameter) opens, just posteromedial to the foramen ovale. The basisphenoid apparently does not contribute to the margin of the foramen ovale or, anteriorly, to the border of the foramen for the optic infundibulum (the combined openings for the foramen rotundum and infra- and superior orbital fissures).

Periotic. The periotic is a structurally complex and phylogenetically informative bone, and deserves detailed description. Figure 11 shows the orientation of the periotic in its original position on the skull. When the periotic is in situ, the ventral foramen of the facial canal can be seen in ventral view, but the fenestra ovalis is partially obscured by the pars cochlearis.

The right periotic ( Figures 12 and 13 ) has a slender anterior process, slightly smaller posterior process, and a dorsoventrally weakly inflated pars cochlearis between these processes. Both the anterior and posterior processes make a wide angle with the anteroposterior axis when the periotic is placed into original position on the skull.

The anterior process is prolonged anteriorly, with a blunt apex. The anterior process is deep and somewhat laterally widened, as seen in Waipatia maerewhenua . Dorsally, a very weak anteroexternal sulcus runs from just anterior to the lateral tuberosity to the anterior end of the dorsal crest. The parabullary sulcus ( Tanaka and Fordyce, 2014) is deep and strongly curved (C-shaped). The anteroventral angle, as seen anteriorly, is a small bump just medial to the anterior bullar facet. There is no obvious anterior keel or anterodorsal angle, and this region of the anterior process is smoothly rounded. A small tubercle on the medial surface of the anterior process has a tiny foramen or vertical canal as seen in W. maerewhenua . On the weak tubercle, a straight groove runs from the anterior incisure. The anterior bullar facet is a shallow groove, about 8.5 mm long, with raised parallel-sided margins (3.7 mm width). The fovea epitubaria is occupied by a slightly damaged and moderate sized accessory ossicle, which is preserved between the anterior bullar facet and the mallear fossa. The ossicle has a thin subrectangular flange on its medial side. The flange has a shallow groove on its margin (see Figures 12 and 13.2 ) and is also widely grooved anteroposteriorly on its dorsal face, possibly forming a sulcus for the tensor tympani muscle. Just posteromedial to the accessory ossicle, there is a tiny foramen. The dorsal surface of the periotic has a prominent dorsal crest, medial to which is a deep anteroposterior groove (9.1 mm), as seen in Otekaikea marplesi but longer and shallower. The anterior incisure is less distinctly grooved than seen in Waipatia maerewhenua and Otekaikea marplesi .

The pars cochlearis is hemispherical ventrally, transversely compressed, and long relative to width. The smooth anteromedial corner passes back into a straight medial margin then an obtuse posteromedial angle. The internal auditory meatus opens on the center medially. A tiny slit-like foramen for the hiatus Fallopii opens on the anterior incisure. The internal auditory meatus is open, posteriorly wider, and rounded tear-shape (maximum length, 8.8 mm). The meatus contains four foramina or groups of foramina: the proximal opening of the facial canal, the foramen singulare, the spiral cribriform tract, and the area cribrosa media. A very low transverse ridge separates the small, narrow proximal opening for the facial canal (2.6 mm) from the foramen singulare, which is also small and narrow. Between the foramen singulare and the spiral cribriform tract, there is a higher crest that bounds a large (anteroposterior length; 5.6 mm), deep and rounded area, which is shared by a cluster of foramina forming the spiral cribriform tract, and a pit for the area cribrosa media. As with some other odontocetes ( Mead and Fordyce, 2009), the area cribrosa media is not clearly perforate. The spiral cribriform tract has a rounded opening and screw structure.

The fenestra rotunda is an anteroposteriorly short compressed reniform opening (transverse diameter 3.3 mm). The aperture for the cochlear aqueduct is subcircular and small (1.3 mm diameter). The small aperture for the vestibular aqueduct is a transversely long ellipse (2.6 mm). Just ventral to the medial margin of the internal auditory meatus, there is a weak groove that might be the median promontorial groove (see Mead and Fordyce, 2009, p. 122).

The posterior extremity of the pars cochlearis is lying lateral to the fenestra rotunda and medial to the fossa for the stapedial muscle, with a small tubercle of the dorsoventrally slightly thin caudal tympanic process, which projects posteroventrally.

On the body of the periotic, ventrally, the mallear fossa is deep, wide, and slightly reniform in shape; the rim of the fossa connects with the accessory ossicle anteriorly. The lateral tuberosity is weakly developed. A short grooved facial crest arises at the base of the mallear fossa and descends posteriorly to the apex of the posterior process. An indistinct depression lateral to the crest at the posterior part of the mallear fossa is probably the fossa incudis. Further posteriorly and lateral to the facial crest, the large hiatus epitympanicus comprises a smooth anterior depression and a rugose grooved posterior part just anterior to and parallel with the face of the posterior process. The narrow groove between the facial crest and the facial sulcus is possibly the tympanohyal sulcus, as seen in Otekaikea marplesi . The fenestra ovalis is rounded, small (2.1 mm), and filled by the footplate of the stapes (the head is missing). The ventral foramen of the facial canal is small, round, and posteriorly directed, located about level with the anterior of the fenestra ovalis; the foramen is slightly covered by the facial crest. The facial sulcus runs along the posteromedial face of the posterior process, close to the posterior bullar facet, becoming indistinct about 3 mm behind the level of the caudal tympanic process (13-14 mm anterior to the tip of the posterior process). The stapedial muscle fossa runs back along the face of the posterior process to within about 10 mm of the tip of the posterior process, separated from the facial sulcus by a faint ridge. Most of the stapedial muscle fossa lies immediately behind the fenestra ovale; the fossa is deep and slightly longer than wide, and forms a subcylindrical depression on the face of the caudal tympanic process.

The posterior process is rectangular and slen- der (17.6 mm long and 7.8 mm wide). The posterior bullar facet has three sections separated by faint ridges: small anterior and posterior sections, and a larger medial section. The dorsal margin of the tip of the process was originally sutured with the squamosal; the periotic was freed by removing matrix with dilute acetic acid and was found to be unfused with the squamosal.

Posterior to the lateral tuberosity, the distinctive articular rim projects laterally. Medial to the articular rim, there are at least three large presumed posteroexternal foramina. A deep groove runs from anteromedial to posterolateral, at the lateral end of the posterior process, so that the anterolateral part of the posterior process is a thin flange. Posterior to the groove, there are many weak striae on the posterolateral end of the posterior process.

Stapes. The stapes has a wide footplate, in situ in the right periotic. The crura are broken at the dorsal level of the stapedial foramen. The footplate is anteroposteriorly long and sub-elliptical, with slightly perpendicular anterior and medial margins, and slightly rounded posterior and lateral margins. A fine crest spans between the two broken crura; whether the stapedial foramen was open is uncertain. Ventrally, the head provides a small articulation for the incus. Just dorsal to the head, posteromedially, there is a large rounded muscular process. This preservation, broken in the midheight, is very similar with the stapes of Prosqualodon davidis ( Flynn, 1948, figure 7), perhaps reflecting weakness caused by a large stapedial foramen. Conversely, some more-crownward platanistoids ( Platanista gangetica and Notocetus vanbenedeni AMNH 29026) reportedly lack the stapedial foramen ( Anderson, 1878; de Muizon, 1987). In Tursiops , the foramen is vestigial and non-patent ( Mead and Fordyce, 2009).

Bulla. Only the rounded left sigmoid process ( Figures 12 and 13 ) is preserved. The dorsal surface lacks the slightly flattened facet that, in some other odontocetes, matches the sigmoid fossa on the squamosal. Anterior to the sigmoid process, on the ventral surface, there is a depressed area with a few weak grooves. In ventral view, the margins of the sigmoid process are thickened. Inside the sigmoid process, the tympanic sulcus runs from the posteromedial end to anterolateral part of the sigmoid process. A broken fossa for malleus is on the medial base of the sigmoid process.

Mandibles. The fused mandibles preserve the symphysis and the alveoli but lack the anterior and posterior ends. The incomplete mandible is 520+ mm long; judging from premaxillary length, at least 170 mm is missing from the anterior.

The anterior part of the mandible is flat ventrally, with at least five mental foramina directed anteriorly on the ventral surface ( Figure 14.7 ). The most anterior preserved foramen is notably long (131 mm) and deep. The anteroposteriorly long and shallow (20.5 mm height) mandibular symphysis has three foramina on its suture on the dorsal surface ( Figure 14.4 ). The damaged part of the symphysis shows internal openings of the mental foramina. Some of the internal areas are filled by cemented matrix, suggesting that the mandible contains canals for the nerves, vessels, and especially the intra-mandibular fat body. Mental foramina are preserved on the ventral surface of the mandible ( Figure 14.8 ). The ventral surface of the mandibular symphysis has a triangular depression between the prominent ventral margins. The dorsal surface is tabular.

Around 28 mm lateral from the posterior end of the mandibular symphysis, the long bilateral alveolar groove opens dorsolaterally, widening posteriorly. Width ranges from 4.5-7.5 mm. Posteriorly, there are slightly elongate alveoli (around 17 mm long and 6 mm width). Here, the body of each mandible swells laterally; the dorsal border rises gradually. The most dorsal part of the coronoid process is broken but the base of the process shows a strong coronoid crest. Medially, each mandibular fossa is long and high. Only the dorsal border is preserved near the coronoid process, indicating a maximum depth of about 95 mm. The profile of the inflated lateral margin suggests that the fossa could have been 150+ mm long.

Teeth. Otekaikea huata is near-homodont and polydont ( Figure 10 ). Fifty-one single-rooted teeth are preserved isolated; in at least one cheek-tooth, the roots are vestigially double but almost completely fused. Tooth positions are identified with reference to other archaic odontocetes, especially Waipatia maerewhenua . The term dimension is used in preference to “length” for overall size, as length is an anteroposterior or mesiodistal measurement, akin to diameter in most of the teeth described below.

An extremely long-rooted tusk ( Figure 10.1-2 ) with a worn crown, presumably an incisor, is thick at the crown and the upper part of the root, tapering and becoming slightly flattened toward the root apex. The rough-surfaced root is long (197+ mm) and becomes thickest (16.1 mm widest diameter) slightly below the crown; this root is around 2.5 times longer and 1.6 times the diameter than the largest tooth in Otekaikea marplesi . The crown is naturally worn, leaving the crown base (11.9 mm maximum diameter at the enamelo-dentin junction) with exposed dentin and a pulp cavity infilled by matrix. Of note, the wear pattern around the crown is asymmetric and elliptical; one side is worn more than the other, retaining a small area of enamel. From another view, the worn end is slightly curved like a shovel ( Figure 10.2 ).

Two more tusks ( Figure 10.3-4 , 93.5 and 93.1 mm, respectively) are shorter than the former tooth. Both are also worn, with exposed dentin. Their diameters are 7.7 mm and 9.6 mm, respectively, narrowing toward the root tip. They have a stronger curve than the large tusk, viewed from the worn surface and are presumed to be posterior incisors (i.e., second or third incisors).

Four conical medium-sized teeth ( Figure 10.5- 6 ) are 48-57 mm in total dimension and 7-8 mm in diameter. In the figured tooth, the crown, which is slightly laterally compressed, comprises 40% of the total dimension; it is slightly worn on the apex. A small enamel slit may indicate the posterior face, judging from the single-rooted teeth of Waipatia maerewhenua ( Tanaka and Fordyce, 2014) . The outer surface (probably labial, rather than buccal) of the conical crown is weakly convex. A little below the crown, the root swells on one face and forms the thickest part on the tooth. Apically (away from the crown), the root becomes thinner and slightly laterally compressed, with a shallow long groove on the presumed labial face.

Thirty-six smaller moderately curved teeth ( Figure 10.7-9 ) are from 31 to 26 mm in total dimension and 5.7-6.7 mm diameter. The crown is about 30% of the total dimension. Loch et al. (2015) examined one tooth from this specimen, and reported the enamel as thin (75-85 microns), with an inner layer of radial enamel and an outer layer of prismless enamel. Some teeth are worn markedly; most show slight wear. They also have a small enamel slit posteriorly.

Eleven buccolingually flattened teeth ( Figure 10.10-12 ) are 21-23 mm total dimension and 6.4- 6.7 mm diameter. Around 30% of the total dimension comprises the crown. There are elongate grooves on the buccal and lingual surfaces of the root.

One tooth, with a large main denticle and two small posterior accessory denticles ( Figure 10.13- 14 ), is the only tooth indicating some vestigial heterodonty, in contrast with the homodont teeth of most extant odontocetes (single rooted teeth with a conical crown and no accessory denticles). Dimensions are 23.2 mm from tip of crown to tip of root, 7.0 mm anteroposteriorly and 5.7 mm buccolingually. The crown comprises 30% of the total dimension. Its anterior keel is worn along a narrow (1.0 mm wide) band. The accessory denticles barely project from the posterior keel; the moreapical (more-anterior) denticle is slightly larger and more prominent than the more basal denticle. Both faces on the crown have subvertical long enamel ridges that are more-regularly subparallel buccally. Lingually, a weak entocingulum is developed. The single root has marked grooves on both buccolingual surfaces, as if marking the junction of two confluent roots. Just below the crown base, the root swells markedly posteriorly.

Atlas. The atlas ( Figure 15.1, 15.6 and 15.11 ) is anteroposteriorly thick and not fused to the axis. The epiphyses are fused with the body. Only basal parts of the neural arch are preserved, including part of the margin of the transverse foramen (for the first spinal nerve and vertebral artery; see Flower, 1885b, figure 15). The neural canal and confluent odontoid fovea have a dorsally wide open V-shape. The maximum dorsal width is 41.9 mm, and height (between a line of both sides of the fragmentary neural arch and the ventral border of the neural canal) is 37.5 mm. The upper transverse process is small (25.0+ mm projection beyond the body) and directed strongly dorsolaterally. The lower process is smaller but more robust than the upper transverse process, is wide (9.2 mm long) and low (10.9 mm in thickness), and projects posterolaterally. The condyloid facets (articular surfaces) for the occipital condyles are deep and distinctly separated ventrally with clear ridges. In lateral view, the condyloid facets face down at about 20 degrees to the plane of the tabular articular facets for the axis. The latter facets are separated by the deep fovea for the odontoid process of the axis. On each articular facet, oblique strong tubercles descend forward to delimit the odontoid region; the transverse ligament (see Struthers, 1872) probably arose from the ridge. From the posteroventral margin of the ventral arch, a long robust hypapophysis (ventral tubercle) projects posteroventrally from an elongate base. The process extends 12.9 mm below and 15.7 mm behind the body. Its cross-section is elliptical.

Axis. The axis ( Figure 15.2, 15.7 , and 15.12) is anteroposteriorly thinner than the atlas and not fused with the adjacent vertebrae. The epiphyses are fused with the body. The neural spine is broken but the transversely robust neural arch curves and rises dorsally. In anterior and posterior view, the neural canal appears ventrally wide and subtriangular, with a prominent median crest that separates bilateral depressions at the base of each pedicle. Each transverse process is thick, high, and strongly projected laterally, with bifurcated extremities. There is no vertebrarterial canal, but the weak depression on the posterior surface of the transverse process is probably the homologous feature. The wide and flat articulations for the atlas anteriorly merge at the smooth surface under the strongly projecting odontoid process. The elliptical (wide) posterior epiphysis is depressed at the center, has a vertical crease, has an indistinct notochordal pit, and has indistinct circumferential striations for the annular ligament. The ventral margin projects, passing into a weak crest on the ventral surface of the body to the odontoid process.

Cervical vertebrae. Three cervical vertebrae ( Figure 15.3-5, 15.7 , and 15.10), probably the fourth to sixth, are preserved; all the vertebral epiphyses are ankylosed. Each vertebra has a tiny neural spine, thin neural arches, a wide rounded-pentagonal neural canal, and strongly ventrolaterally projecting and wide transverse processes, each inferred to have formed a vertebrarterial canal. The anteroposteriorly flattened body has a slightly convex anterior face, and has a sagittal ventral keel and a low dorsal sagittal crest separating bilateral dorsal depressions. On the lateral surface of the body, there is a deep and large depression, dorsal to the transverse process. The bodies of the posterior cervical vertebrae are anteroposteriorly thicker than the most anterior one.

Thoracic vertebrae. Twelve thoracic vertebrae are preserved ( Figure 16 ; not all are figured), all with the anterior epiphyses ankylosed. Each thoracic vertebra has: a high, long, and flat neural spine; a small fovea (synovial articulation) for the rib on the robust transverse process, and a larger fovea on the body; a weakly depressed anterior surface of the body; an anteroposteriorly long body (lengthens progressively posteriorly in the series); and no distinctive ridge on ventral surface of the body. On the dorsal surface of the body, there are paired anteroposteriorly long oval dorsal foramina.

Thoracic vertebra 1 on Figure 16 is assumed to be an anterior thoracic vertebra because the transverse process arises in a lower position. It has a wide and low neural canal, short body and anteroposteriorly shortest neural arch. Two to four thoracic vertebrae are more posterior and have an anteroposteriorly long neural spine, high pentagonal shaped neural canal, and a longer body. The transverse process arises from a high point dorsally on the neural arch. Vertebra 4 shows a short and blunt transverse process from the dorsal part of the body. It has a deep and tear-shaped fossa on the anterior base of the transverse process.

Two lumbar vertebrae ( Figure 16.6 ) have the anterior epiphyses ankylosed. Each lumbar vertebra has a very long wing-like transverse process; flat anterior and posterior surfaces of the body; and an anteroposteriorly long body. Vertebra F on Figure 16 is 139.3+ mm wide across the transverse processes and 54.4+ mm long, while another lumbar vertebra (not pictured) is 52.5 mm wide and 43.1 mm long.

Ribs. In total, 12 ribs ( Figure 17); two right and two left, and six others with only the mid-part of the shaft (their sides are uncertain) are preserved. Two right and left anterior double-headed ribs, damaged ventrally, are 180+ and 176+ mm chord length, respectively, anteroposteriorly flat and wide. There is a diffuse triangular and shallow cavity on the posterior surface. On the shaft, anterior and posterior surfaces are weakly swollen and flat, respectively. The rib portion converges distally.

Four presumed mid-series of ribs show a conical shaft with slightly anteroposterior flattening, and with medial and lateral blunt margins. The longest preserved rib is 402+ mm chord length, around 13 mm length and 20 mm width around at the mid-shaft. The anterior and posterior surfaces are weakly swollen and flat, respectively. A left rib has the strong tubercle at around 70 mm distal to the capitulum on the lateral edge. Two right ribs show that both dorsal and ventral ends are widened dramatically, twice as wide as the mid-shaft. The dorsal end is short (or anteroposteriorly flat). The ventral end is thick and shows transversely long elliptical surface for the cartilaginous sternal ribs.

Sternum. The symmetrical flat manubrium (or presternum) ( Figure 18 ) has incomplete outlines. Dimensions are 96.9+ mm long, 85.6+ mm wide, and 17.4 mm thick. The presumed ventral side is ventrally warped. The presumed anterior is wider, perhaps providing an area for the cartilaginous sternal ribs.

Scapula. The right scapula is better preserved ( Figures 19 , 20 ), but the left scapula ( Figure 20 ) retains the coracoid process, which is broken on the right. For this description, the orientation of the scapula has the glenoid fossa ventral, and the suprascapular border dorsal as figured ( Figure 19 ). The fan-shaped scapula is transversely thin and longer than high and has two projections (the acromion and coracoid) anteriorly.

The scapular blade is anteroposteriorly long (wide in lateral view), with the angle of the lines from the anterior and posterior borders is 106°. Anteriorly, the narrow prescapular (supraspinous) fossa and the large postscapular (infraspinous) fossa are separated by a rounded ridge. Both fossae have a smooth undulating surface; there is strong depression at the center of the scapula. The posterior convexity for the border between the infraspinatus and the teres major lies slightly posterior to the center of the scapula. On the medial surface, two ridges may be origins for tendons of the subscapularis muscle.

The acromion projects slightly ventrally when the glenoid cavity of the scapula is sitting on a flat plane, and curves gently laterally at the base and stops anteromedially. It provides a huge space for the supraspinatus muscle. The long (94 mm) acromion is parallel-sided anteriorly. A relatively small rod-like coracoid is long relative to diameter (25.3 mm long, versus diameters of 8.3 mm and 7.8 mm at mid-length), and robust at the base (13.7 mm width, 14.2 mm height). The neck of the scapula is strongly constricted. The scapula has a deep rounded glenoid cavity ventrally with a small foramen in the center.

Humerus. The right and left humeri are well preserved ( Figures 19.2 and 20.5-8 ). The humerus is long, cylindrical (153 mm), and transversely slightly flattened (40 mm width and 65.4 mm length at the mid-shaft). For identifications of the muscle attachments, our direct examination of Globicephala melas and previous studies on the Odontoceti are used ( Schulte and Smith, 1918; Howell, 1930; Smith et al., 1976; Strickler, 1978).

Proximally, a rounded, transversely long, elliptical, and smooth head is exposed laterally, occupying around 30% of the total length of the humerus. Medial to the head, there is a wide flat area for the subscapularis muscle. Below the head, there are weak anterior and posterior constrictions (proximal view). The small tuberosity lies posterior to the head and provides a small corner probably for the triceps muscle. Between the tuberosities, the bicipital groove appears to be absent. Anterior to the angle for the coracobrachialis, there is a large steep step for the supraspinatus muscle, formed by the proximal part of the humeral anterior margin. Anterior to the head, distal to the steep step for the supraspinatus, is a large proximodistally long depression for the infraspinatus muscle, beyond which is a large nutrient foramen at the level of the mid-shaft. The anterior margin of the humerus is thin and sigmoidal. Just slightly proximal to the fossa for the infraspinatus, there is a rough surface for the mastohumeralis muscle on the anterior edge. At the same level with the fossa for the infraspinatus, there is a small tubercle, probably for the serratus muscle on the posterior margin of the humerus. Just proximal to the distal end of the humerus is a weak crescent depression for the deltoid muscle. On the medial surface, muscle origins are uncertain (i.e., the teres major and minor). Distally, the joints for the ulna and radius are smooth with weak depressions on their centers. These joints are separated by a ridge. The posterior angle carries the joint for the ulna and has a small tubercle anteriorly and a strongly projected tubercle (an attachment for the triceps muscle) posteriorly. The anterior half of the distal end of the humerus is the joint for the radius.

Ulna. The right ulna is robust proximally and transversely thin distally ( Figures 19.3 and 20.10 ). The olecranon projects posteriorly and is slightly curved laterally from proximal and distal view. The base of the olecranon and the proximal articular face for the humerus are thick (or wide). The trochlear notch is strongly curved (angle around 90°). The interosseous and the outer borders are straight, and gradually and weakly widened distally. On the distal end, the epiphysis is not fused. The distal end is an anteroposteriorly long ellipse (length 47.9+ mm and width 22.8+ mm).

Radius. The right and left radii ( Figure 19.4 ) are preserved. The right radius shows an exact fit with the humerus. The shaft is weakly curved anteriorly and slightly thinner than the ulna. Both proximal and distal epiphyses are not fused. The proximal end is triangular with a small projection at the anterior end of the posteriorly-tilted articular face for the humerus. The anterior margin is thin, and the posterior margin shows a flattened area. The shaft widens gradually to the distal end.

Carpus. A single carpus ( Figure 20.9 ) is 40+ mm long, 26 mm wide, and 18.3 mm thick. It may be a proximal carpus because it is very wide and is almost the same width as the ulna and radius.

Figures

FIGURE 1. Locality map and stratigraphic sections of the Otekaikea huata type locality. Arrow shows type locality. Foraminiferal zonation and geochronology are based on Graham et al. (2000).

FIGURE 2. Dorsal views of the type skull, OU 22306, Otekaikea huata (1 and 2). 3, dorsolateral view, figure is not to scale.

FIGURE 3. Ventral views of the type skull, OU 22306, Otekaikea huata. Pa, the parietal.

FIGURE 4. Anterior views of the type skull, OU 22306, Otekaikea huata.

FIGURE 5. Posterior views of the type skull, OU 22306, Otekaikea huata.

FIGURE 6. Right lateral views of the type skull, OU 22306, Otekaikea huata.

FIGURE 7. Disarticulated rostral elements, OU 22306, Otekaikea huata. 1, dorsal view. 2, left premaxilla lateral view. 3, right premaxilla lateral view. 4, ventral view. 5 and 6, detailed photos of the premaxilla.

Figure 8. Skull of Otekaikea huata, OU 22306. Right 1, lateral view and 2, dorsal view.

FIGURE 9. Details of right basicranium part, OU 22306, Otekaikea huata. 1 and 2, ventral view. 3 and 4, ventrolateral view. Ali, alisphenoid. Bo, basioccipital. Bs, basisphenoid. Pa, parietal. Vo, vomer.

FIGURE 10. The type teeth, OU 22306, Otekaikea huata. Scales vary as indicated, depending on specimen size. 1 and 2, largest incisor. 1, worn surface view. 2, lateral view. 3 and 4, two long tusks, showing worn surface. 5 and 6, two medium teeth. 7-9, single rooted smaller medium teeth. 10-12, buccolingually flattened teeth. 13 and 14, tooth with small denticles. 13, buccal view. 14, lingual view.

FIGURE 11. Periotic in original position in the right squamosal, ventral view, OU 22306, Otekaikea huata.

FIGURE 12. The type tympanoperiotics. OU 22306, Otekaikea huata. 1-6, right periotic. 7 and 8, sigmoid process of left bulla. 1, medial view. 2, lateral view. 3, ventral view. 4, dorsal view. 5, posterior view. 6, anterior view. 7, ventral view. 8, posterior view.

FIGURE 13. Key features of the type tympanoperiotics. OU 22306, Otekaikea huata. 1-6, right periotic. 7 and 8, sigmoid process of left bulla. 1, medial view. 2, lateral view. 3, ventral view. 4, dorsal view. 5, posterior view. 6, anterior view. 7, ventral view. 8, posterior view.

FIGURE 14. The type mandible, OU 22306, Otekaikea huata. 1, anterior view. 2, left lateral view. 3, dorsal view. 4, mandibular symphysis in dorsal view. 5, posterior view. 6, ventral view. 7, anterior end in ventral view. 8, right lateral view.

FIGURE 15. The type cervical vertebrae, OU 22306, Otekaikea huata. 1-5, anterior views. 6-8, lateral views. 9-10, dorsal views. 1, atlas. 2 and 8, axis, 3, fourth cervical vertebra. 4, fifth cervical vertebra. 5, sixth cervical vertebra. 6 and 9, atlas and axis. 7 and 10, third to sixth cervical vertebrae. 7, mirrored image.

FIGURE 16. The type thoracic and lumbar vertebrae, OU 22306, Otekaikea huata. 1-3, anterior view of three thoracic vertebrae. 4 and 5, a posterior thoracic vertebra, anterior and left lateral views. 6, lumbar vertebrae from anterior view.

FIGURE 18. The type sternum. OU 22306, Otekaikea huata. 1, ventral. 2, left lateral. 3, dorsal.

FIGURE 19. The type right forelimb bones in lateral view, OU 22306, Otekaikea huata. 1, right scapula. 2, humerus. 3, ulna. 4, radius.

FIGURE 20. The type forelimb bones, OU 22306, Otekaikea huata. 1, right scapula in medial view. 2, right scapula in distal view. 3, left scapula in lateral view. 4, left scapula in distal view. 5-8, right humerus. 5, posterior view. 6, medial view. 7, anterior view. 8, proximal view. 9, carpus. 10, ulna in posterior view.