Arktocara yakataga Boersma &amp; Pyenson, 2016, sp. nov.
Creators: name: Boersma, Alexandra T.
name: Pyenson, Nicholas D.
Publisher: name: Zenodo
Year: 2016
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Mammalia
Cetacea
Allodelphinidae
Arktocara
Arktocara yakataga

description: Arktocara yakataga, sp. nov. (Figs. 2 10 and Table 1) LSID: urn:lsid:zoobank.org:act: FBCF 0EAA­7BBB­4EF0­8186­7548993098D1 Holotype. USNM 214830, consisting of an incomplete skull lacking the rostrum anterior of the antorbital notches, tympanoperiotics, dentition and mandibles (see Fig. 2). Type locality. The precise geographic coordinates for the type locality of Arktocara yakataga are unknown. The type specimen (USNM 214830) was discovered and collected in 1951 by United States Geological Survey (USGS) geologist Donald J. Miller (1919 1961), who was mapping what was then the Yakataga District of Alaska (now the Yakutat City and Borough) between 1944 and 1963. Archival notes housed with the specimen at USNM state that Miller found the specimen in the Poul Creek Formation within the then­Yakataga District (see Age, below). Therefore, we delimit the area for the type’s provenance to exposures of the Poul Creek Formation in the Yakutat City and Borough, Alaska, USA, in a grid ranging approximately from 60 22 0N, 142 30 0W to 60 0 0 0N, 143 22 0W (see Fig. 1). While the formation has been named from its exposures along Poul Creek, it has been suggested that the most abundant macrofossils from this unit have been collected from outcrops along Hamilton Creek, White River, and Big River near Reare Glacier (Taliaferro, 1932). It is possible that Miller collected USNM 214830 from one of these exposures. Formation. Poul Creek Formation. Age. Archival documentation accessioned in the Department of Paleobiology with USNM 214830 indicate that the type specimen was collected from an unknown locality exposed about 400 500 m below the top of the Poul Creek Formation, which has a total stratigraphic thickness of around 1.9 km (Plafker, 1987). The Yakutat terrane of Southeast Alaska consists of the Kulthieth, Poul Creek, and Yakataga Formations (Perry, Garver &amp; Ridgway, 2009; Plafker, Moore &amp; Winkler, 1994; Miller, 1971). The Kulthieth Formation consists of mostly organic­rich sandstones deposited in nonmarine alluvial, deltaic, barrier beach and shallow marine environments, and is Early Eocene to Early Oligocene (54 33 Ma) in age based on the fossil assemblages present (Perry, Garver &amp; Ridgway, 2009). The Upper Eocene to possibly Lower Miocene (40 20 Ma) Poul Creek Formation conformably overlies the Kulthieth Formation (Plafker, 1987; Miller, 1971). It is estimated to be approximately 1.9 km thick, and is composed of siltstones and organic­rich sandstones, in part glauconitic recording a marine transgression, interrupted by deposits of the Cenotaph Volcanics (Plafker, 1987). Finally, unconformably overlying the Poul Creek Formation is the Miocene to Pliocene Yakataga Formation (Miller, 1971). It is composed mainly of tillite and marine strata (Perry, Garver &amp; Ridgway, 2009). The Poul Creek Formation itself is broadly constrained to approximately 40 20 million years in age, from the latest Eocene to possibly early Miocene in age (Plafker, 1987; Miller, 1971). The depositional age of the unit has been further constrained to 24 to 29 Ma, or a mid to late Oligocene age, based on detrital zircon fission­track analyses of young grain­age populations (Perry, Garver &amp; Ridgway, 2009). Using the broadest time duration for the formation (20 million years) and the coarse stratigraphic thickness of the sediments within it (2 km), a constant rate of sedimentation would suggest that the stratigraphic position of USNM 214830 at 500 m below the top of the formation would be roughly equivalent to an geochronologic age of 25 million years, an estimate that is consistent to the detrital zircon analyses. Overall, we propose a late Oligocene, or Chattian age for Arktocara, although we cannot exclude a Rupelian antiquity. Diagnosis. Arktocara is a small to medium sized platanistoid odontocete (approximately 2.26 m in total length), which belongs, based on one equivocal synapomorphy, to the node­based Platanistoidea: width: width of the premaxillae&gt;50% of the width of the rostrum at the antorbital notch (character 51[1]). More convincingly, Arktocara belongs to Platanistoidea based on its affinities to other members of the Allodelphinidae that possess unequivocal synapomorphies of the Platanistoidea (see ‘Discussion’ for further comments on the relationship of Allodelphinidae within the Platanistoidea). We also note that, for the purposes of this diagnosis, we use a broad definition of Waipatiidae that includes Otekaikea spp. (see Tanaka &amp; Fordyce (2015a)), and Squalodelphinidae sensu Lambert, Bianucci &amp; Urbina (2014). See ‘Discussion’ for further comments on systematics of these groups. Arktocara can be differentiated from all other platanistoids by the following combination of character states. Arktocara differs from Waipatiidae and Squalodelphinidae in having: a postglenoid process of squamosal greatly reduced (character 114[1]); an occipital shield bearing a distinct sagittal crest (character 118[1]); length of the zygomatic process as percent of the greatest width of the maxillae across the postorbital processes &lt;30% (character 152[1]); lacking a dorsal condyloid fossa (character 119[0]); and lacking any asymmetry in the vertex (character 98[0]). Arktocara also differs from Waipatiidae and Platanistidae in having: frontals posterior to the nasals and between the premaxillae wider than the maximum transverse width across the nasals (character 95[0]); and lacking an anterior transverse ridge and large tympanosquamosal recess, with middle sinus still inferred to be present (character 144[1]). Arktocara further differs from Waipatiidae in having: a lacrimal that wraps around the anterior edge of the supraorbital process of frontal and slightly overlies its anterior end (character 37[0]); maxilla forming the dorsolateral edge of the internal opening of the infraorbital foramen (character 43[0]); ventromedial edge of the internal opening of infraorbital foramen formed by maxilla and palatine (character 44[1]); a nuchal crest weakly convex anteriorly in dorsoposterior view (character 117[1]); a lateral end of the groove for the mandibular branch of the trigeminal nerve wrapping laterally around posterior end of pterygoid sinus fossa and opening primarily anteriorly (character 148[0]); the angle formed by the basioccipital crests in ventral view between 15 40 (character 157[1]); and in lacking a premaxillary crest or posterior maxillary crest adjacent to the nasal (character 72[0]). Arktocara differs from all Platanistidae and Squalodelphinidae in having the anterolateral corner of the maxilla overlying the supraorbital process of frontal being thin and of even thickness to parts posteromedial (character 64[0]). Arktocara also differs from all Platanistidae in having: the apex of the postorbital process of frontal projected posterolaterally and slightly ventrally (character 46[0]); the ratio of the greatest width of the premaxillae to the greatest width of maxillae at the level of postorbital processes between 0.49 0.38 (character 76[1]); a shallow emargination of the posterior edge of zygomatic process by the sternomastoid muscle fossa in lateral view (character 111[1]); the width of the squamosal lateral to the exoccipital in posterior view as a percent of the greatest width of the exoccipitals &lt;15% (character 112[0]); fossae for the preorbital lobe of the pterygoid sinus present in the orbit (character 134[1]); shallow posterior portion of the periotic fossa of the squamosal (character 151[1]); posteroventral­most point on the basioccipital crest forming a closely appressed flange with a narrow crease separating exoccipital dorsally from the rest of crest (character 156[1]); and lacking a pneumatic maxillary crest overhanging medially (character 65[0]). Finally, Arktocara differs from all Squalodelphinidae in having: a narrower width of the premaxillae at the antorbital notches as a percent width of the rostrum at the antorbital notch (50 64%) (character 51[1]). Arktocara differs from all other Allodelphinidae in having: a reduced postglenoid process of the squamosal (character 114[1]); and the posteroventral­most point of the basioccipital crest forming a closely appressed flange separating the exoccipital dorsally from the rest of the crest by a narrow crease (character 156[1]). Arktocara differs from Allodelphis pratti and Goedertius oregonensis in having: both premaxillae extending posterior to the nasals (character 58[1]); and the ratio of the greatest width of the premaxillae to greatest width of the maxillae at the level of the postorbital processes between 0.49 0.38 (character 76[1]). Arktocara also differs from Goedertius oregonensis and Zarhinocetus errabundus in having: the width of the premaxillae between 50 64% of the width of the maxillae a the level of the antorbital notches (character 51[1]); a uniformly thin anterolateral corner of the maxilla overlying the supraorbital process of the frontal (character 64[0]); length of the zygomatic process of the squamosal 30% of the width of the maxillae at the postorbital processes (character 152[1]); lacking a rostral basin (character 50[0]), lacking a posterior dorsal infraorbital foramen placed posteromedially near the posterior extremity of the premaxilla (character 60[0]); lacking a dorsal condyloid fossa (character 119[0]) and having a symmetrical cranial vertex (character 98[0]). Arktocara also differs from Allodelphis pratti and Zarhinocetus errabundus in having: two anterior dorsal infraorbital foramina (character 49[1]); and a shallow emargination of the posterior edge of the zygomatic process by the sternomastoid muscle fossa in lateral view (character 111[1]). Arktocara further differs from Goedertius oregonensis having: a fused lacrimal and jugal (character 39[1]); the apex of the postorbital process of the frontal directed posterolaterally and slightly ventrally (character 46[0]); a triangular or anteroposteriorly widened falciform shaped postorbital process of the frontal (character 47[2]); one posterior dorsal infraorbital foramina of the maxilla (character 59[1]); nuchal crest weakly convex anteriorly in dorsoposterior view (character 117[1]); and the posterior edge of the vomer terminating on the basisphenoid (character 154[0]). Arktocara further differs from Zarhinocetus errabundus in having: the transverse width of the nasal&gt;70% the length of the nasal (character 91[2]); a distinct sagittal crest on the occipital shield (character 118[1]); and in lacking an anterior transverse ridge and large tympanosquamosal recess (character 144[1]). Lastly, Arktocara displays the following apomorphies: a weakly developed antorbital notch (character 10[0]); straight lateral margin of the right premaxilla posterior to premaxillary foramen (character 56[1]); a U­shaped nasal frontal suture (character 94[2]); and a greatly reduced postglenoid process of the squamosal (character 114[1]). Etymology. The species epithet ‘yakataga’ derives from the Tlingit name for the point of land along the southeast coast of Alaska between modern day Kayak Island and Ice Bay. This point, currently called Cape Yakataga, is located directly southwest of Watson Peak and represents the southeastern boundary of a floodplain drained by the Bering Glacier. The name Yakataga was first published by Tebenkov (1852: map 7), who was a cartographer and hydrographer of the Imperial Russian Navy, as ‘‘M[ys] Yaktaga’’ on an 1849 map of Alaska. The geographic place name has been alternatively spelled Cape Iaktag, Cape Yakaio, Cape Yakatag, and Yokataga Reef (Orth, 1967). According to the Geographic Names Information System (GNIS, 2016), developed by USGS in cooperation with the United States Board of Geographic Names (BGN), the name ‘‘Yakataga’’ means ‘‘canoe road,’’ referring to two reefs that form a canoe passage to the shore of the village. Description Anatomical terminology follows Mead &amp; Fordyce (2009). In most cases, description of individual elements derives from the most informative side of the skull, in terms of preservation; we note any morphological asymmetry if present. Besides overall erosion of the bone surfaces and loss of some skull elements, there appears to be no significant burial­related distortion of the skull. <i>Skull</i> The holotype of Arktocara yakataga (USNM 214830) consists of an incomplete skull, measuring 23 cm in preserved length. The majority of the rostrum is missing, with an asymmetric transverse break approximately 2 5 cm anterior to the antorbital notch. The skull also lacks both nasals, jugals, tympanoperiotics, and the right occipital condyle (see Figs. 2 and 3). Most of the paroccipital processes of the exoccipitals are gone, large portions of the palatines and pterygoids are missing, small fragments along the lateral margins of the frontals and maxillae are incomplete, and the general condition of many osteological elements in the skull are poorly preserved. The skull may have been both mechanically and chemically prepared in the past (with no known documentation), including acid preparation, which may have contributed to the poor state of preservation for the osteological surfaces of many elements. Portions of the skull are obscured by a fine­grained grey matrix of siltstone, especially infilling the mesorostral canal, the bony nares, the recesses of tympanoperiotic region, and the braincase (which is exposed via the foramen magnum and fenestrae in the supraoccipital). In dorsal view, the preserved skull is roughly hexagonal in overall shape (Fig. 2). The external nares are vertically oriented, and positioned at a level between the antorbital and postorbital processes. The vertex is particularly table­like and square, composed of frontals, premaxillae, and nasals (missing). The parietals appear to be narrowly exposed on the vertex immediately anterior to the nuchal crest, between the frontals and supraoccipital whether this is natural, or an ontogenetic characteristic, is unclear. The vertex is bordered anteriorly by the externals nares, laterally by the maxillae and posteriorly by the nuchal crest of the supraoccipital. The nuchal crest is straight along the posterior edge of the vertex, but begins to curve posterolaterally as it approaches the temporal crest. The temporal fossae are visible in dorsal view due to an intertemporal constriction just anterior to the level of the nuchal crest, and the fossa is floored by a narrow valley (the squamosal fossa) between the squamosal plate and supramastoid crest. In lateral view, the profile of the skull gradually slopes upwards from the level of the antorbital notch to posterior of the nares, where it levels out on the vertex (Fig. 4). The nuchal crest is well defined, and about the same height as the frontals on the vertex. The orbit is shallowly rounded dorsally (4.9 cm in length), with the maxilla completely overlying the frontal on the thin supraorbital process, except on the postorbital process, where the frontal is exposed laterally. It is unclear whether the antorbital process of the frontal is completely covered by maxilla or not, but most of the medial antorbital process is composed of the lacrimojugal. The temporal fossa is trapezoidal in shape, with the temporal crest forming a right angle with the dorsal margin of the zygomatic process of the squamosal. The dorsal margin of the temporal fossa is roofed over by the frontal. The preserved posterior portion of rostrum anterior of the antorbital notch is wide (8.8 cm anterior to the antorbital notch) and deep (6.5 cm at the level of the antorbital notch), with a broadly open and deep mesorostral canal (2.4 cm wide and 4.6 cm deep at the level of the antorbital notch). While there is material missing around the antorbital notch, the posterior margin of the right antorbital notch appears to be real, demonstrating it to be weakly developed, forming an L­shape with the lateral margin of the rostrum (Fig. 2). Though some of the anteromedial antorbital process is missing, not enough is missing to have drastically changed the shape of the antorbital notch region. In anterior view, the maxilla abruptly slopes upwards medially to the distinct premaxilla­maxilla suture, and underlies the premaxilla along the entire preserved length of rostrum and likely the cranium (Fig. 5). The premaxilla therefore forms an anteroposteriorly elongated rectangular plateau surrounding the external bony nares, elevated in relation to the plane of the maxillae, appearing similar in transverse cross­section to a horst and graben system. In ventral view, there is clearly a fossa for the preorbital lobe of the pterygoid sinus (a shallow depression surrounding the ventral infraorbital and sphenopalatine foramina), as well as for the hamular lobe of the pterygoid sinus (fossa anterior to the choanae). Also present are an extremely shallow tympanosquamosal recess for the middle sinus, and a middle pterygoid sinus fossa anterior to the periotic fossa (Fig. 6). There is not enough rostral material or paroccipital process to determine the presence of an anterior or posterior sinus respectively, and the presence of a peribullary sinus is difficult to ascertain. <i>Premaxilla</i> Both of the premaxillae are symmetrical, and overlie either the maxillae or the frontals for their entire preserved length (Fig. 2). In lateral view, the premaxilla thins slightly as it passes around the external nares, otherwise maintaining a relatively even thickness on the cranium (Fig. 4). The premaxilla also appears to thin anterior of the antorbital notch (especially in right lateral view), lowering to the same level as the maxilla instead of rising dorsally above it. However, in anterior view, it is evident that the ventral margin of the left premaxilla sinks ventrally into a medial trough created by the maxilla while the dorsal margin remains level, accounting for the apparent reduction in thickness (Fig. 5). Both premaxillae are broken anteriorly, the right further anterior than the left and missing some of its anterolateral edge. The premaxilla­maxilla suture is clear in dorsal view along the entire lateral length of the premaxilla, as well as in anterior view at the transverse cross­section of the rostrum. In dorsal view, the lateral margin of the premaxilla is mostly rectilinear, widening only 0.5 cm from the rostral break to a level anterior to the nares. As it passes laterally around the nares, the premaxilla gently bows out laterally, with the medial edge retreating more than the lateral edge so that the total width is reduced (0.8 cm on the right premaxilla). Posterior to the nares, the lateral edge remains straight posteriorly, but the medial edge expands slightly medially, once again widening the premaxilla. On the left side of the skull, lateral and posterior of the external nares, a narrow ledge of the medial margin of the maxilla laterally borders the premaxilla, where the premaxilla is separated from the maxilla (possibly diagenetically). On the vertex, the posterior termination of the premaxilla lies on the frontal as an asymmetrical, spatulate lobe, tapering posterolaterally and bordered medially by frontal and the nasal fossa of the frontal, and laterally the maxilla (Fig. 2). There is no division of the premaxilla into a posteromedial splint and posterolateral plate (a widespread characteristic amongst odontocetes). The premaxillary sac fossa, roughly at the level of the antorbital notch, is shallow with a rough surface. No premaxillary foramina or associated premaxillary su
description: Published as part of <i>Alexandra T. Boersma &amp; Nicholas D. Pyenson, 2016, Arktocara yakataga, a new fossil odontocete (Mammalia, Cetacea) from the Oligocene of Alaska and the antiquity of Platanistoidea, pp. 1-41 in Peerj 2321</i> on pages 9-25, DOI: 10.7717/peerj.2321, http://zenodo.org/record/270212
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