Leonard Dewaele

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.

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.

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.

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.

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.

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.

Generalized Osteosclerotic Condition in the Skeleton of Nanophoca vitulinoides , a Dwarf Seal from the Miocene of Belgium

In the fossil record, it has been shown that various clades of secondarily aquatic tetrapods experienced an initial densification of their bones in the early stages of their evolution, and developed spongier and lighter bones only later in their evolution, with the acquisition of more efficient swimming modes. Although the inner bone structure of most secondarily aquatic tetrapods has already been studied, no research hitherto focused on true seals, or Phocidae. However, preliminary observations previously made on a Miocene species, Nanophoca vitulinoides, suggested that this taxon showed pronounced specialization of bone structure as compared to other seals. This feature justifies a specific comparative study, which is the purpose of this article. Microanatomical analysis of bones of N. vitulinoides shows compactness values nearing 100%, which is much higher than in other semi-aquatic mammals, pinnipeds included. Osteohistological analyses show virtually complete remodeling of the medullary territory by Haversian substitution. Extreme bone compactness locally resulted from an imbalance, towards reconstruction, of this process. Cortical regions were less intensely remodeled. In a number of specimens, the cortex shows clear growth marks as seasonal lines of arrested growth. The results suggest that, despite the extreme compactness of long bones of N. vitulinoides and the small size of this taxon, the growth rate of the cortex, and that of the bones in general, did not differ strongly from that of other, larger phocids. Extreme skeletal compaction and densification must have increased body density in Nanophoca. Consequently, speed, acceleration, and maneuverability must have been low, and this taxon was most likely a near-shore bottom-dwelling seal. Consequently, dietary preferences were most likely oriented towards benthic food sources.