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Arktocara yakataga
Boersma & Pyenson, 2016
Artistic reconstruction of a pod of Arktocara yakataga, swimming offshore of Alaska during the Oligocene, about 25 million years ago, with early mountains of Southeast Alaska in the background. The authors speculate that Arktocara may have socialized in pods, like today's oceanic dolphins, while possessing a much longer snout, like its closest living relative in the freshwater rivers of South Asia.
Linocut print art by Alexandra Boersma |
Abstract
The diversification of crown cetacean lineages (i.e., crown Odontoceti and crown Mysticeti) occurred throughout the Oligocene, but it remains an ongoing challenge to resolve the phylogenetic pattern of their origins, especially with respect to stem lineages. One extant monotypic lineage, Platanista gangetica (the Ganges and Indus river dolphin), is the sole surviving member of the broader group Platanistoidea, with many fossil relatives that range from Oligocene to Miocene in age. Curiously, the highly threatened Platanista is restricted today to freshwater river systems of South Asia, yet nearly all fossil platanistoids are known globally from marine rocks, suggesting a marine ancestry for this group. In recent years, studies on the phylogenetic relationships in Platanistoidea have reached a general consensus about the membership of different sub-clades and putative extinct groups, although the position of some platanistoid groups (e.g., Waipatiidae) has been contested. Here we describe a new genus and species of fossil platanistoid, Arktocara yakataga, gen. et sp. nov. from the Oligocene of Alaska, USA. The type and only known specimen was collected from the marine Poul Creek Formation, a unit known to include Oligocene strata, exposed in the Yakutat City and Borough of Southeast Alaska. In our phylogenetic analysis of stem and node-based Platanistoidea, Arktocara falls within the node-based sub-clade Allodelphinidae as the sister taxon to Allodelphis pratti. With a geochronologic age between ∼29–24 million years old, Arktocara is among the oldest crown Odontoceti, reinforcing the long-standing view that the diversification for crown lineages must have occurred no later than the early Oligocene.
Systematic paleontology
Cetacea Brisson, 1762
Odontoceti Flower, 1867 sensu Fordyce & Muizon, 2001
Platanistoidea (CCN) (node-based version of Fordyce, 1994)
Allodelphinidae (CCN) (node-based version of Barnes, 2006)
Arktocara, gen. nov.
LSID: urn:lsid:zoobank.org:act:EE11B95B-8338-496B-97F4-1673ED90E709
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The skull of Arktocara yakataga on an 1875 ethnographic map of Alaska drawn by William Healey Dall, a broadly trained naturalist who worked for several US government agencies, including the Smithsonian, and honored with several species of living mammals, including Dall's porpoise (Phocoenoides dalli). Near the skull of Arktocara is a cetacean tooth, likely belonging to a killer whale (Orcinus orca), collected by Aleš Hrdlička, a Smithsonian anthropologist who worked extensively in Alaska, and an Oligocene whale tooth collected by Donald Miller, a geologist who worked for the US Geological Survey, and collected the type specimen of Arktocara. Donald Orth's dictionary of Alaskan place names, published by the USGS, bookends the image.
photo: James Di Loreto, Smithsonian |
Definitions. Crown group Platanista refers to the crown clade arising from the last common ancestor of all lineages descending from Platanista, including two subspecies of Platanista gangetica (P. g. gangetica (Lebeck, 1801) and P. g. minor Owen, 1853), as recognized by The Society for Marine Mammology’ Committee on Taxonomy (2015).
Type and only included species: Arktocara yakataga, sp. nov.
Etymology. The name Arktocara derives from the combination of arktos from Greek and cara from Latin, which together signify “the face of the North.” The only preserved material of the type specimen, USNM 214830 consists of the cranium, or its face, and its type locality is the furthest north that a platanistoid has ever been found.
Age. Same as that of the species.
Diagnosis. Same as that of the species.
Arktocara yakataga, sp. nov. (Figs. 2–10 and Table 1)
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The skull of Akrtocara yakataga rests on an 1875 ethnographic map of Alaska drawn by William Healey Dall, a broadly trained naturalist who worked for several US government agencies, including the Smithsonian, and honored with several species of living mammals, including Dall's porpoise (Phocoenoides dalli). Near the skull of Arktocara is a cetacean tooth, likely belonging to a killer whale (Orcinus orca), collected by Aleš Hrdlička, a Smithsonian anthropologist who worked extensively in Alaska, and an Oligocene whale tooth collected by Donald Miller, a geologist who worked for the US Geological Survey, and collected the type specimen of Arktocara. Donald Orth's dictionary of Alaskan place names, published by the USGS, bookends the image.
photo: James Di Loreto, Smithsonian |
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 then-Yakataga 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′N, 142°30′W to 60°00′N, 143°22′W (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 & Ridgway, 2009; Plafker, Moore & Winkler, 1994; Miller, 1971). The Kulthieth Formation consists of mostly organic-rich 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 & 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 organic-rich 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 & 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 fission-track analyses of young grain-age populations (Perry, Garver & 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 node-based Platanistoidea: width: width of the premaxillae >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 & Fordyce (2015a)), and Squalodelphinidae sensu Lambert, Bianucci & Urbina (2014). See ‘Discussion’ for further comments on systematics of these groups.
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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.
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Figure 12: Distribution map of fossil Allodelphinidae.
Mapped of fossil localities of allodelphinids, projected on a truncated Winkel Tripel map and centered on 25°N and 170°W. Occurrences for fossil data derive from location of type and referred localities for each taxon, are listed alphabetically by region, and are represented by orange dots.
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Platanistoids first appear in the fossil record in the late Oligocene, and reach peak richness in the early Miocene (Kimura & Barnes, 2016; Tanaka & Fordyce, 2015a). The oldest platanistoids with solid age constraints are the waipatiids, all found in the Oligocene-Miocene Otekaike Limestone of New Zealand (Graham et al., 2000; Benham, 1935; Fordyce, 1994; Tanaka & Fordyce, 2014; Tanaka & Fordyce, 2015a). Based on both the lithology and the presence of age-diagnostic planktic foraminifera and ostracod species, Waipatia hectori (Benham, 1935) is the oldest reported waipatiid, from the uppermost Duntroonian Stage of the Otekaike Limestone, approximately 25.2 Ma (Tanaka & Fordyce, 2015b). Arktocara is possibly very similar in age to Waipatia hectori, constrained to the Chattian Stage of the upper Oligocene in the Poul Creek Formation, approximately ∼24–29 Ma (Perry, Garver & Ridgway, 2009). Unfortunately, the lack of robust locality data for either Waipatia hectori or Arktocara makes impossible to determine which is the oldest.
Arktocara is, however, very clearly the oldest known allodelphinid, expanding the previously reported age range of Allodelphinidae by as much as 9 million years (Kimura & Barnes, 2016). Other allodelphinids span temporally from the early to middle Miocene, which largely matches the stratigraphic range of other platanistoid lineages (Fig. 11). Interestingly, Arktocara is among the oldest crown Odontoceti, reinforcing the long-standing view that the timing for the diversification for crown lineages must have occurred no later than the early Oligocene.
Lastly, Allodelphinidae appear uniquely limited, in terms of geography, to marine rocks of the North Pacific Ocean, with occurrences in Japan, Alaska, Washington State, Oregon, and California (see Fig. 12; Kimura & Barnes, 2016). Arktocara expands this geographic range to sub-Arctic latitudes. At approximately 60°N in the Yakutat City and Borough, Arktocara is the most northern platanistoid yet reported. The next most northern platanistoid reported is an incomplete and unnamed specimen from the late Chattian marine Vejle Fjord Formation in northern Denmark, approximately 56.7°N, 9.0°E (Hoch, 2000).
Alexandra T. Boersma and Nicholas D. Pyenson. 2016.
Arktocara yakataga, a new fossil odontocete (Mammalia, Cetacea) from the Oligocene of Alaska and the antiquity of Platanistoidea.
PeerJ. 4:e2321. DOI:
10.7717/peerj.2321
New species of extinct river dolphin discovered in Smithsonian collection
bit.ly/2aGgRS8 via @EurekAlertAAAS
