Robert W. Boessenecker

Anatomy, feeding ecology, and ontogeny of a transitional baleen whale: a new genus and species of Eomysticetidae (Mammalia: Cetacea) from the Oligocene of New Zealand

The Eocene history of cetacean evolution is now represented by the expansive fossil record of archaeocetes elucidating major morphofunctional shifts relating to the land to sea transition, but the change from archaeocetes to modern cetaceans is poorly established. New fossil material of the recently recognized family Eomysticetidae from the upper Oligocene Otekaike Limestone includes a new genus and species, Waharoa ruwhenua, represented by skulls and partial skeletons of an adult, juvenile, and a smaller juvenile. Ontogenetic status is confirmed by osteohistology of ribs. Waharoa ruwhenua is characterized by an elongate and narrow rostrum which retains vestigial alveoli and alveolar grooves. Palatal foramina and sulci are present only on the posterior half of the palate. The nasals are elongate, and the bony nares are positioned far anteriorly. Enormous temporal fossae are present adjacent to an elongate and narrow intertemporal region with a sharp sagittal crest. The earbones are characterized by retaining inner and outer posterior pedicles, lacking fused posterior processes, and retaining a separate accessory ossicle. Phylogenetic analysis supports inclusion of Waharoa ruwhenua within a monophyletic Eomysticetidae as the earliest diverging clade of toothless mysticetes. This eomysticetid clade also included Eomysticetus whitmorei, Micromysticetus rothauseni, Tohoraata raekohao, Tokarahia kauaeroa, Tokarahia lophocephalus, and Yamatocetus canaliculatus. Detailed study of ontogenetic change demonstrates postnatal elaboration of the sagittal and nuchal crests, elongation of the intertemporal region, inflation of the zygomatic processes, and an extreme proportional increase in rostral length. Tympanic bullae are nearly full sized during early postnatal ontogeny indicating precocial development of auditory structures, but do increase slightly in size. Positive allometry of the rostrum suggests an ontogenetic change in feeding ecology, from neonatal suckling to a more specialized adult feeding behaviour. Possible absence of baleen anteriorly, a delicate temporomandibular joint with probable synovial capsule, non-laterally deflected coronoid process, and anteroposteriorly expanded palate suggests skim feeding as likely mode of adult feeding for zooplankton. Isotopic data in concert with preservation of young juveniles suggests the continental shelf of Zealandia was an important calving ground for latitudinally migrating Oligocene baleen whales.

New Records of the Fur Seal Callorhinus (Carnivora: Otariidae) from the Plio-Pleistocene Rio Dell Formation of Northern California and Comments on Otariid Dental Evolution

ABSTRACT New fossils representing two species of the fur seal Callorhinus are reported from the uppermost Pliocene to lower Pleistocene Rio Dell Formation of northern California. The finds include latest Pliocene-earliest Pleistocene dentarles and postcrania of Callorhinus gilmorei, and a partial dentary of early Pleistocene age identified as Callorhinus sp. The aforementioned material is ascribed to C. gilmorei due to the incipient single-rooted condition of the p1–2, retention of double-rooted p3–m1, and overall small size. The dentary identified as Callorhinus sp. exhibits a more derived pattern of tooth morphology, including single-rooted p1–p4 (and double-rooted ml), larger size than C. gilmorei, and in the size range of extant Callorhinus ursinus (which typically exhibit fused roots on all postcanine teeth). Fusion of postcanine roots began with the p2 and continued posteriorly, and is likely an adaptation to accommodate crowded teeth anteriorly in the jaws. Callorhinus gilmorei has previously been reported from the upper Pliocene of southern California and Japan, and this new record extends the range of this taxon further north in the Northeast Pacific. Callorhinus sp. is the most complete pinniped fossil to be described from the early Pleistocene of the Northeast Pacific. The wide biogeographic range of Callorhinus during the Pliocene and Pleistocene documents the persistence of this taxon, potentially as a Pliocene-Holocene anagenetic lineage. This highlights the antiquity of the Callorhinus lineage, which has persisted in the Northeast Pacific since the Pliocene, establishing it as the oldest and earliest diverging crown otariid.

Comparative Taphonomy, Taphofacies, and Bonebeds of the Mio-Pliocene Purisima Formation, Central California: Strong Physical Control on Marine Vertebrate Preservation in Shallow Marine Settings

Background Taphonomic study of marine vertebrate remains has traditionally focused on single skeletons, lagerstätten, or bonebed genesis with few attempts to document environmental gradients in preservation. As such, establishment of a concrete taphonomic model for shallow marine vertebrate assemblages is lacking. The Neogene Purisima Formation of Northern California, a richly fossiliferous unit recording nearshore to offshore depositional settings, offers a unique opportunity to examine preservational trends across these settings. Methodology/Principal Findings Lithofacies analysis was conducted to place vertebrate fossils within a hydrodynamic and depositional environmental context. Taphonomic data including abrasion, fragmentation, phosphatization, articulation, polish, and biogenic bone modification were recorded for over 1000 vertebrate fossils of sharks, bony fish, birds, pinnipeds, odontocetes, mysticetes, sirenians, and land mammals. These data were used to compare both preservation of multiple taxa within a single lithofacies and preservation of individual taxa across lithofacies to document environmental gradients in preservation. Differential preservation between taxa indicates strong preservational bias within the Purisima Formation. Varying levels of abrasion, fragmentation, phosphatization, and articulation are strongly correlative with physical processes of sediment transport and sedimentation rate. Preservational characteristics were used to delineate four taphofacies corresponding to inner, middle, and outer shelf settings, and bonebeds. Application of sequence stratigraphic methods shows that bonebeds mark major stratigraphic discontinuities, while packages of rock between discontinuities consistently exhibit onshore-offshore changes in taphofacies. Conclusions/Significance Changes in vertebrate preservation and bonebed character between lithofacies closely correspond to onshore-offshore changes in depositional setting, indicating that the dominant control of preservation is exerted by physical processes. The strong physical control on marine vertebrate preservation and preservational bias within the Purisima Formation has implications for paleoecologic and paleobiologic studies of marine vertebrates. Evidence of preservational bias among marine vertebrates suggests that careful consideration of taphonomic overprint must be undertaken before meaningful paleoecologic interpretations of shallow marine vertebrates is attempted.

Pleistocene survival of an archaic dwarf baleen whale (Mysticeti: Cetotheriidae)

Pliocene baleen whale assemblages are characterized by a mix of early records of extant mysticetes, extinct genera within modern families, and late surviving members of the extinct family Cetotheriidae. Although Pleistocene baleen whales are poorly known, thus far they include only fossils of extant genera, indicating Late Pliocene extinctions of numerous mysticetes alongside other marine mammals. Here a new fossil of the Late Neogene cetotheriid mysticete Herpetocetus is reported from the Lower to Middle Pleistocene Falor Formation of Northern California. This find demonstrates that at least one archaic mysticete survived well into the Quaternary Period, indicating a recent loss of a unique niche and a more complex pattern of Plio–Pleistocene faunal overturn for marine mammals than has been previously acknowledged. This discovery also lends indirect support to the hypothesis that the pygmy right whale (Caperea marginata) is an extant cetotheriid, as it documents another cetotheriid nearly surviving to modern times.

A Reevaluation of the Morphology, Paleoecology, and Phylogenetic Relationships of the Enigmatic Walrus Pelagiarctos

Background A number of aberrant walruses (Odobenidae) have been described from the Neogene of the North Pacific, including specialized suction-feeding and generalist fish-eating taxa. At least one of these fossil walruses has been hypothesized to have been a specialized predator of other marine mammals, the middle Miocene walrus Pelagiarctos thomasi from the Sharktooth Hill Bonebed of California (16.1–14.5 Ma). Methodology/Principal Findings A new specimen of Pelagiarctos from the middle Miocene “Topanga” Formation of southern California (17.5–15 Ma) allows a reassessment of the morphology and feeding ecology of this extinct walrus. The mandibles of this new specimen are robust with large canines, bulbous premolars with prominent paraconid, metaconid, hypoconid cusps, crenulated lingual cingula with small talonid basins, M2 present, double-rooted P3–M1, single-rooted P1 and M2, and a P2 with a bilobate root. Because this specimen lacks a fused mandibular symphysis like Pelagiarctos thomasi, it is instead referred to Pelagiarctos sp. This specimen is more informative than the fragmentary holotype of Pelagiarctos thomasi, permitting Pelagiarctos to be included within a phylogenetic analysis for the first time. Analysis of a matrix composed of 90 cranial, dental, mandibular and postcranial characters indicates that Pelagiarctos is an early diverging walrus and sister to the late Miocene walrus Imagotaria downsi. We reevaluate the evidence for a macropredatory lifestyle for Pelagiarctos, and we find no evidence of specialization towards a macrophagous diet, suggesting that Pelagiarctos was a generalist feeder with the ability to feed on large prey. Conclusions/Significance This new specimen of Pelagiarctos adds to the knowledge of this problematic taxon. The phylogenetic analysis conclusively demonstrates that Pelagiarctos is an early diverging walrus. Pelagiarctos does not show morphological specializations associated with macrophagy, and was likely a generalist predator, feeding on fish, invertebrates, and the occasional warm-blooded prey item.

Unique feeding morphology in a new prognathous extinct porpoise from the Pliocene of California.

Modern porpoises (Odontoceti: Phocoenidae) are some of the smallest cetaceans and usually feed near the seafloor on small fish and cephalopods [1-3]. Within both extinct and extant phocoenids, no evidence for specialized mandibular morphology has been documented [4-7]. Here we describe a new species of extinct porpoise, Semirostrum ceruttii, from the marine Pliocene San Diego (4.2-1.6 mega-annum, Ma) and Purisima (5-2.5 Ma) formations of California. The mandibles comprise a long, fused, and nearly edentulous prognathous symphysis, extending farther beyond the rostrum than in any known mammal. Phylogenetic analyses based on morphology reconstruct Semirostrum ceruttii as sister to extant (crown) porpoise species with moderate support. We describe the spectacularly preserved holotype specimen based on computed tomography (CT) scans, which allowed visualization of the elongate mental and accessory canals within the symphysis. The elongate canals are similar to those found in Rynchops birds [8] and were likely involved in sensory function. Oblique labial wear facets present on numerous small conical mandibular teeth posterior to the symphysis suggest regular contact with benthic substrate. The unique mandibular and dental characteristics, along with robust scapulae, sternum, and unfused cervical vertebrae, support the interpretation that this species employed a form of benthic skim feeding by using its mandible to probe for and obtain prey.

Latest Pacific Basin record of a bony-toothed bird (aves, Pelagornithidae) from the Pliocene Purisima Formation of California, U.S.A.

ABSTRACT Fossils of pelagornithids (bony-toothed birds) have been reported from strata of Paleocene to Pliocene age, and from every continent. The extreme fragility of pelagornithid bones has no doubt contributed to their geographically and temporally sporadic record, and thus it has been difficult to appreciate any long-term phylogenetic trends through geologic time for this group. We report a well-preserved partial humerus of the gigantic bird Pelagornis from the late Neogene Purisima Formation of central California. Due to its incompleteness, we refrain from naming a new species. This fossil is fortuitously bracketed by two ash beds, which have been correlated with volcanic rocks at 3.35 ± 0.05 Ma and 2.5 ± 0.2 Ma, indicating a middle to late Pliocene age for this fossil. This fossil extends the record of the pelagornithids in the Northeast Pacific, previously only known up until the early late Miocene (10–12 Ma). This fossil is the latest record of a pelagornithid for the Pacific Basin, and additionally represents the latest reliably dated pelagornithid record worldwide. This record suggests that the pelagornithids survived until the end of the Pliocene, and became extinct during the ocean restructuring and climatic upheavals that caused the demise of many other groups of marine vertebrates at that time.

A new eomysticetid from the Oligocene Kokoamu Greensand of New Zealand and a review of the Eomysticetidae (Mammalia, Cetacea)

Eomysticetids represent a worldwide, short-lived radiation of archaic baleen-bearing mysticetes that elucidate morphofunctional shifts and ontogenetic change amongst early Neoceti. Fossils of eomysticetids are now recorded from Oligocene marine rocks along the margins of the North Pacific (Japan), North Atlantic (South Carolina), and the Southern Ocean (New Zealand). The assemblage from New Zealand is diverse, with five species in three genera present during the Duntroonian stage (Chattian, 27.3–25.2 Ma). A new genus and species of eomysticetid from the lower Kokoamu Greensand of New Zealand, Matapa waihao gen. et sp. nov., increases the known diversity of Southern Hemisphere eomysticetids. Matapa uniquely exhibits a rounded margin of the occipital shield, and shares a mosaic of features seen in Northern Hemisphere eomysticetids such as a large squamosal prominence, an inflated and posteriorly rounded paroccipital process, and a double-faced posterior bullar facet with longitudinal ridges. The holotype specimen of Matapa waihao is ontogenetically immature, but ontogenetically static tympanoperiotic characters that diagnose this new species are shared by referred adult specimens. Matapa is uppermost Whaingaroan (28.1–27.3 Ma) in age, the oldest eomysticetid from New Zealand. Fossils of Matapa are recovered from both the modern west and east coast of New Zealand which, during the Oligocene, were on the margin of the Southern Ocean. Inclusion of Matapa within an earlier published cladistic analysis recovers Matapa as the earliest diverging member of the New Zealand eomysticetid clade, and confirms eomysticetid monophyly. The taxonomy, morphology and geochronological age of all known eomysticetids are reviewed. http://zoobank.org/urn:lsid:zoobank.org:pub:EE1CB5CC-618B-4ABA-9D97-5B9DEDD3470E

Mammalian bite marks on juvenile fur seal bones from the late Neogene Purisima Formation of central California

Abstract Fossils of extinct fur seals and walruses (Carnivora: Pinnipedia) occur within rich vertebrate fossil assemblages recovered from the shallow marine Mio-Pliocene Purisima Formation, central California. Two isolated postcranial bones—a humerus and a radius—belonging to a juvenile fur seal (Pinnipedia: Otariidae) exhibit circular depressions. These bone modifications are associated with radial and circular fractures, and are characterized by inward displacement of the cortex. These depressions lack features typical of erosive invertebrate borings, trampling damage from media ( = substrate) interaction, puncturing by another object during diagenetic compaction, such as a clast embedded or associated with the modification, or pathologic bone modification. These features are best interpreted as tooth marks. These tooth marks lack certain characteristics of commonly reported marks inflicted by shark teeth, such as linear gouges and subparallel scrapes formed by xiphodont and serrated teeth. These bone modifications instead exhibit a circular shape and inward displacement of the cortex, consistent with puncturing by a conical mammal tooth. The size and distribution of the tooth marks, in concert with the known vertebrate assemblage from the Purisima Formation, indicate several possible producers of the bone modifications: a pilot whale or beluga-like cetacean, a terrestrial carnivore, a dusignathine or odobenine walrus, or a case of infanticide by a conspecific otariid.

The oldest known fur seal

The poorly known fossil record of fur seals and sea lions (Otariidae) does not reflect their current diversity and widespread abundance. This limited fossil record contrasts with the more complete fossil records of other pinnipeds such as walruses (Odobenidae). The oldest known otariids appear 5–6 Ma after the earliest odobenids, and the remarkably derived craniodental morphology of otariids offers few clues to their early evolutionary history and phylogenetic affinities among pinnipeds. We report a new otariid, Eotaria crypta, from the lower middle Miocene ‘Topanga’ Formation (15–17.1 Ma) of southern California, represented by a partial mandible with well-preserved dentition. Eotaria crypta is geochronologically intermediate between ‘enaliarctine’ stem pinnipedimorphs (16.6–27 Ma) and previously described otariid fossils (7.3–12.5 Ma), as well as morphologically intermediate by retaining an M2 and a reduced M1 metaconid cusp and lacking P2–4 metaconid cusps. Eotaria crypta eliminates the otariid ghost lineage and confirms that otariids evolved from an ‘enaliarctine’-like ancestor.