Sexual dimorphism in the walrus mandible: comparative description and geometric morphometrics.

The modern walrus Odobenus rosmarus is characterized by marked sexual dimorphism, related to its polygynous behavior and the aggressive competition between males during the breeding season. Previous studies treated skeletal sexual dimorphism in walruses either qualitatively or with basic quantitative measurements. The present study combines a detailed qualitative comparison of male and female walrus mandibles with quantitative two-dimensional geometric morphometrics analysis (principal component analysis, Procrustes ANOVA and a linear discriminant analysis). In addition to identifying previously recognized sexually dimorphic features (e.g., convexity of the anterior margin of the mandible in adult males), our study finds new morphological differences between males and females, such as a relative dorsal expansion of the anterior part of the mandible and an accentuated concavity between the dorsal margin and the coronoid process in adult males. Both our qualitative comparisons and quantitative analyses demonstrate that sexual dimorphism as expressed in the mandible of extant walruses is statistically significant and that (variation in) mandibular morphology can be used as tool to attribute sex with a good degree of accuracy to isolated mandibles or skeletons lacking the cranium. Sexual dimorphism in walruses is directly related to their sexual behavior, characterized as aggressive in males and linked to a polygynous reproduction system. Indeed, the difference in size of the tusks between males and females but also the use of these during intraspecific fights, can reasonably account for this great mandibular morphological disparity between adult males and females, but also among different ontogenetic stages. Finally, the results obtained in the present study may serve as a starting point for assessing sexual dimorphism more in-depth and studying inter- and intraspecific variation in the mandibles of fossil walruses by identifying quantified size and shape mandibular features.

Hypersalinity drives convergent bone mass increases in Miocene marine mammals from the Paratethys.

Pachyosteosclerosis-a condition that creates dense, bulky bones-often characterizes the early evolution of secondarily aquatic tetrapods like whales and dolphins1-3 but then usually fades away as swimming efficiency increases.4 Here, we document a remarkable reversal of this pattern, namely the convergent re-emergence of bone densification in Miocene seals, dolphins, and whales from the epicontinental Paratethys Sea of eastern Europe and central Asia. This phenomenon was driven by imbalanced remodeling and inhibited resorption of primary trabeculae and coincided with hypersaline conditions-the Badenian salinity crisis-that affected the Central Paratethys between 13.8 and 13.4 Ma.5 Dense bones acting as ballast would have facilitated efficient swimming in the denser and more buoyant water and hence were likely adaptive in this setting. From the Central Paratethys, pachyosteosclerosis subsequently spread eastward, where it became a defining feature of the endemic late Miocene whale assemblage.6,7.

A late surviving Pliocene seal from high latitudes of the North Atlantic realm: the latest monachine seal on the southern margin of the North Sea.

BACKGROUND: The family of true seals, the Phocidae, is subdivided into two subfamilies: the southern Monachinae, and the northern Phocinae, following the subfamilies' current distribution: extant Monachinae are largely restricted to the (sub-)Antarctic and the eastern Pacific, with historical distributions of the monk seals of the genus Monachus in the Caribbean, the Mediterranean and around Hawaii; and Phocinae to the northern temperate and Arctic zones. However, the fossil record shows that Monachinae were common in the North Atlantic realm during the late Miocene and early Pliocene. Until now, only one late Pliocene record is known from the Mediterranean, Pliophoca etrusca from Tuscany, Italy, but none from farther north in the North Atlantic. METHODS: We present the description of one partial phocid humerus collected in the early 20th century from the Antwerp area (Belgium), with an assessment of its stratigraphic origin using data from the literature. RESULTS: The studied humerus was recovered during construction works at the former Lefèvre dock in the Antwerp harbour (currently part of the America dock). Combining the information associated to the specimen with data from the literature and from local boreholes, the upper Pliocene Lillo Formation is ascertained as the lithological unit from which the specimen originates. Morphologically, among other features the shape of the deltopectoral crest and the poor development of the supinator crest indicates a monachine attribution for this specimen. The development of the deltopectoral crest is closer to the condition in extant Monachinae than in extinct Monachinae. DISCUSSION: The presented specimen most likely represents a monachine seal and a literature study clearly shows that it came from the latest early to late Pliocene Lillo Formation. This would be the first known monachine specimen from the latest early to late Pliocene of the North Sea, and more broadly from the northern part of the North Atlantic realm. This humerus differs from the humerus of P. etrusca and suggests a higher diversity of Monachinae in the latest early to late Pliocene than previously assumed.

A critical revision of the fossil record, stratigraphy and diversity of the Neogene seal genus Monotherium (Carnivora, Phocidae).

Historically, Monotherium had been one of the few genera of extinct Phocidae (true seals) that served as a wastebin taxon. Consequently, it did neither aid in understanding phylogenetic relationships of extinct Phocidae, nor in understanding seal diversity in deep time. This urged the reassessment of the genus. Before our review, Monotherium included five different species: Monotherium aberratum, Monotherium affine, and Monotherium delognii from Belgium; Monotherium gaudini from Italy; and Monotherium? wymani from the east coast USA. In this work we redescribe the fossil record of the genus, retaining the type species M. delognii. Monotherium aberratum and M. affine are reassigned to the new phocine genus Frisiphoca. Monotherium gaudini is renamed and considered a stem-monachine (Noriphoca gaudini). The holotype of the monachine M.? wymani requires further study pending the discovery of new fossil material that could be attributed to the same taxon. Reinvestigating the stratigraphic context reveals that N. gaudini most likely represents one of the two oldest named phocid seals, or even the oldest, dated to the late Oligocene-earliest Miocene. Our results allow questioning the widespread idea that Phocidae originated in the western Atlantic and better appreciate their palaeobiogeography during the late Oligocene-Miocene interval in the North Atlantic realm.

Diversity of late Neogene Monachinae (Carnivora, Phocidae) from the North Atlantic, with the description of two new species.

While the diversity of 'southern seals', or Monachinae, in the North Atlantic realm is currently limited to the Mediterranean monk seal, Monachus monachus, their diversity was much higher during the late Miocene and Pliocene. Although the fossil record of Monachinae from the North Atlantic is mainly composed of isolated specimens, many taxa have been erected on the basis of fragmentary and incomparable specimens. The humerus is commonly considered the most diagnostic postcranial bone. The research presented in this study limits the selection of type specimens for different fossil Monachinae to humeri and questions fossil taxa that have other types of bones as type specimens, such as for Terranectes parvus. In addition, it is essential that the humeri selected as type specimens are (almost) complete. This questions the validity of partial humeri selected as type specimens, such as for Terranectes magnus. This study revises Callophoca obscura, Homiphoca capensis and Pliophoca etrusca, all purportedly known from the Lee Creek Mine, Aurora, North Carolina, in addition to their respective type localities in Belgium, South Africa and Italy, respectively. C. obscura is retained as a monachine seal taxon that lived both on the east coast of North America and in the North Sea Basin. However, H. capensis from North America cannot be identified beyond the genus level, and specimens previously assigned to Pl. etrusca from North America clearly belong to different taxa. Indeed, we also present new material and describe two new genera of late Miocene and Pliocene Monachinae from the east coast of North America: Auroraphoca atlantica nov. gen. et nov. sp., and Virginiaphoca magurai nov. gen. et nov. sp. This suggests less faunal interchange of late Neogene Monachinae between the east and west coasts of the North Atlantic than previously expected.

Reappraisal of the extinct seal "Phoca" vitulinoides from the Neogene of the North Sea Basin, with bearing on its geological age, phylogenetic affinities, and locomotion.

BACKGROUND: Discovered on the southern margin of the North Sea Basin, "Phoca" vitulinoides represents one of the best-known extinct species of Phocidae. However, little attention has been given to the species ever since its original 19th century description. Newly discovered material, including the most complete specimen of fossil Phocidae from the North Sea Basin, prompted the redescription of the species. Also, the type material of "Phoca" vitulinoides is lost. METHODS: "Phoca" vitulinoides is redescribed. Its phylogenetic position among Phocinae is assessed through phylogenetic analysis. Dinoflagellate cyst biostratigraphy is used to determine and reassess the geological age of the species. Myological descriptions of extant taxa are used to infer muscle attachments, and basic comparative anatomy of the gross morphology and biomechanics are applied to reconstruct locomotion. RESULTS: Detailed redescription of "Phoca" vitulinoides indicates relatively little affinities with the genus Phoca, but rather asks for the establishment of a new genus: Nanophoca gen. nov. Hence, "Phoca" vitulinoides is recombined into Nanophoca vitulinoides. This reassignment is confirmed by the phylogenetic analysis, grouping the genus Nanophoca and other extinct phocine taxa as stem phocines. Biostratigraphy and lithostratigraphy expand the known stratigraphic range of N. vitulinoides from the late Langhian to the late Serravallian. The osteological anatomy of N. vitulinoides indicates a relatively strong development of muscles used for fore flipper propulsion and increased flexibility for the hind flipper. DISCUSSION: The extended stratigraphic range of N. vitulinoides into the middle Miocene confirms relatively early diversification of Phocinae in the North Atlantic. Morphological features on the fore- and hindlimb of the species point toward an increased use of the fore flipper and greater flexibility of the hind flipper as compared to extant Phocinae, clearly indicating less derived locomotor strategies in this Miocene phocine species. Estimations of the overall body size indicate that N. vitulinoides is much smaller than Pusa, the smallest extant genus of Phocinae (and Phocidae), and than most extinct phocines.

On Prophoca and Leptophoca (Pinnipedia, Phocidae) from the Miocene of the North Atlantic realm: redescription, phylogenetic affinities and paleobiogeographic implications.

BACKGROUND: Prophoca and Leptophoca represent the oldest known genera of phocine seals, dating from the latest early to middle Miocene. Originally, Prophoca rousseaui and Prophoca proxima were described based on fragmentary remains from the Miocene of Belgium. However, several researchers contested the union of Prophoca rousseaui and Prophoca proxima into one genus, without providing evidence. The stratigraphic context of Prophoca remained poorly constrained due to the lack of precise data associated with the original specimens collected in the area of Antwerp (north of Belgium). METHODS: Prophoca and Leptophoca are redescribed and their phylogenetic position among Phocidae is reassessed using PAUP. Dinoflagellate biostratigraphy has been carried out on sediment samples associated with specimens from Prophoca and Leptophoca to elucidate their approximate ages. RESULTS: Whereas the species Prophoca rousseaui is redescribed, Prophoca proxima is considered synonymous to Leptophoca lenis, with the proposal of a new combination Leptophoca proxima (Van Beneden, 1877). Sediment samples from specimens of both taxa have been dated to the late Langhian-early Serravallian (middle Miocene). Following a reinvestigation of Leptophoca amphiatlantica, characters from the original diagnosis are questioned and the specimens of Leptophoca amphiatlantica are considered Leptophoca cf. L. proxima. In a phylogenetic analysis, Prophoca rousseaui and Leptophoca proxima constitute early branching stem-phocines. DISCUSSION: Leptophoca proxima from the North Sea Basin is younger than the oldest known find of Leptophoca proxima from North America, which does not contradict the hypothesis that Phocinae originated along the east coast of North America during the late early Miocene, followed by dispersal to Europe shortly after. Morphological features of the appendicular skeleton indicate that Prophoca rousseaui and Leptophoca proxima have archaic locomotory modes, retaining a more prominent use of the fore flipper for aquatic propulsion than extant Phocidae.

Perinatal Specimens of Saurolophus angustirostris (Dinosauria: Hadrosauridae), from the Upper Cretaceous of Mongolia.

BACKGROUND: The Late Cretaceous Nemegt Formation, Gobi Desert, Mongolia has already yielded abundant and complete skeletons of the hadrosaur Saurolophus angustirostris, from half-grown to adult individuals. METHODOLOGY/PRINCIPAL FINDINGS: Herein we describe perinatal specimens of Saurolophus angustirostris, associated with fragmentary eggshell fragments. The skull length of these babies is around 5% that of the largest known S. angustirostris specimens, so these specimens document the earliest development stages of this giant hadrosaur and bridge a large hiatus in our knowledge of the ontogeny of S. angustirostris. CONCLUSIONS/SIGNIFICANCE: The studied specimens are likely part of a nest originally located on a riverbank point bar. The perinatal specimens were buried by sediment carried by the river current presumably during the wet summer season. Perinatal bones already displayed diagnostic characters for Saurolophus angustirostris, including premaxillae with a strongly reflected oral margin and upturned premaxillary body in lateral aspect. The absence of a supracranial crest and unfused halves of the cervical neural arches characterize the earliest stages in the ontogeny of S. angustirostris. The eggshell fragments associated with the perinatal individuals can be referred to the Spheroolithus oogenus and closely resemble those found in older formations (e.g. Barun Goyot Fm in Mongolia) or associated with more basal hadrosauroids (Bactrosaurus-Gilmoreosaurus in the Iren Dabasu Fm, Inner Mongolia, China). This observation suggests that the egg microstructure was similar in basal hadrosauroids and more advanced saurolophines. COMPETING INTERESTS: One of the authors (FE) is employed by the commercial organization Eldonia. Eldonia provided support in the form of a salary for FE, but did not have any additional role or influence in the study design, data collection and analysis, decision to publish, or preparation of the manuscript and it does not alter the authors' adherence to all the PLoS ONE policies on sharing data and materials.