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Although modern porpoises (Phocoenidae, Odontoceti, Cetacea) occupy most oceanic regions of the world, their fossil record outside the Pacific Ocean remains extremely scarce. Based on a partial skull from lower Pliocene layers (Kattendijk Formation, Zanclean, 5–4.4 Ma) of the Antwerp harbour, a new genus and species of fossil phocoenid, Brabocetus gigaseorum gen. et sp. nov., is described. The new taxon is diagnosed by a unique combination of characters including an anterolateraly widely-open antorbital notch, the anteromedial sulcus being shorter than the posteromedial sulcus, the frontal boss being longer than wide, pronounced asymmetry of the vertex, and the possible presence of an additional longitudinal sulcus across the premaxillary eminence. The phylogenetic analysis indicates that B. gigaseorum is not closely related to extant phocoenids; it is included in a clade comprising most north-west Pacific fossil species, two north-east Pacific fossil species, and the only previously described North Sea fossil species Septemtriocetus bosselaersi. Considering the paleobiogeographic distribution of phocoenids, an early Pliocene arrival from the North Pacific to the North Sea may be correlated to the early opening of the Bering Strait and migration via the Arctic. The presence of S. bosselaersi in younger deposits (Piacenzian) of the southern margin of the North Sea Basin could be the result of either a second dispersal event, or of local speciation in the North Sea, whereas an additional trans-Arctic migration from the North Pacific lead to the present occupation of the North Atlantic and North Sea by the phylogenetically distantly related harbor porpoise Phocoena phocoena.
In 1991, Sabath described “larger avian eggs” from the Upper Cretaceous Barun Goyot and Djadokhta Formations of Mongolia. These were later included in the ootaxon Gobioolithus major. Here we recognize the larger avian eggs of Sabath as a distinct ootaxon, Styloolithus sabathi, oogen. et oosp. nov. These eggs differ from those of Gobioolithus in being larger (70 by 32 mm) and more elongate. Microscopically, the shell bears a third layer (possible external zone) thicker than the mammillary layer and nearly as thick as the second layer (possible squamatic zone); the continuous layer (including layers two and three) to mammillary layer thickness ratio is 3.1:1. Within the clutch, the tightly spaced eggs stand with their long axes steeply inclined. Adult remains are associated with two clutches, suggesting an incubation mode similar to that of troodontid maniraptorans, where adults sat atop largely buried eggs. S. sabathi provides evidence that relative egg size in Mesozoic non-ornithuromorph birds had increased markedly from the non-avian theropod condition in oviraptorids and troodontids, but had not yet reached the modern egg-adult proportions of Neornithes. Sediment-bound upright eggs appear common to Enantiornithes and more basal avians, suggesting that like non-avian theropods, these birds lacked chalazae, the chords of albumen allowing egg rotation in modern birds. Absence of this simple structure may have restricted these basal birds to ground nesting in areas with appropriate substrates and not permitted the type of nesting diversity found in Neornithes. Neornithes are the only Mesozoic clade of Dinosauria to nest completely free of sediment; this may have played a crucial role in their surviving the K-Pg mass extinction event.
The origin and phylogenetic relationships of most modern coleoid groups have not yet been explained by reliable fossil evidence, in large part because of the reduction or disappearance of a calcified chambered shell during their evolutionary history. Herein we describe two exceptionally large coleoid lower jaws from the Upper Cretaceous strata in Hokkaido, Japan. On the basis of the comparison of gross morphology and morphometric data of the lower jaws of modern and fossil coleoids, we assigned the two lower jaws to the following new taxa: Nanaimoteuthis hikidai sp. nov. of the order Vampyromorpha (superorder Octobrachia) and Haboroteuthis poseidon gen. et sp. nov. of the order Teuthida (superorder Decabrachia). The lower jaw of N. hikidai is distinguished from other species of the same genus from the Upper Cretaceous of Vancouver Island (Canada) and Hokkaido by having a broader, more anteriorly curved hood of the outer lamella. The lower jaw of H. poseidon seemingly exhibits mosaic features like those of modern teuthids and sepiids but is assigned to Teuthida on the basis of the overall shape of the outer lamella and the development of a distinct fold on the lateral wall. Because of the unusually large lower jaws, these new taxa appear to be comparable in body size to modern giant squids (Architeuthis spp.) and the Humboldt squid (Dosidicus gigas). This and other discoveries of large jaws referable to octobrachian and decabrachian coleoids from the Upper Cretaceous strata of the North Pacific fill the gap in the relatively poor fossil record of mainly soft-bodied coleoids.
During the early Palaeozoic, echinoderm body plans were much more diverse than they are today, displaying four distinct types of symmetry. This included the bilateral ctenocystoids, which were long thought to be restricted to the Cambrian. Here, we describe a new species of ctenocystoid from the Upper Ordovician of Scotland (Conollia sporranoides sp. nov.). This allows us to revise the genus Conollia, which was previously based on a single poorly-known species from the Upper Ordovician of Wales (Conollia staffordi). Both these species are characterized by a unique morphology consisting of an elongate-ovoid body covered in spines, which clearly distinguishes them from their better-known Cambrian relatives; they are interpreted as infaunal or semi-infaunal burrowers from deep-water environments. This indicates that the ctenocystoid body plan was not fixed early in the evolution of the group, and they most likely modified their structure as an adaptation to a new mode of life in the Ordovician.
We describe new fossil bovid craniodental remains from the Upper Miocene fossil site of Şerefköy-2, Yatağan Basin, SW Turkey. The new material belongs to six species: Gazella cf. G. capricornis, Palaeoryx pallasi, Sporadotragus parvidens, Skoufotragus cf. Sk. schlosseri, Urmiatherium rugosifrons, and ?Sinotragus sp., which together indicate a latest middle—early Late Turolian (Late Miocene) age. Medium-to-large bovid taxa prevail over small ones, and protoryxoid bovids clearly dominate the assemblage. An analysis of the taxonomic structure, size and diet spectra of several Turolian bovid assemblages from Greece and Turkey reveals Şerefköy-2 to be a member of a mammalian palaeocommunity particular to southwestern Anatolia, which in turn forms part of the sub-Paratethyan biogeographic province.
We describe the first bovid fossils from the late middle Miocene (13.4–13.2 Ma) of the Mae Moh Basin of Northern Thailand, and assign the material to the new species Eotragus lampangensis sp. nov., Eotragus cf. lampangensis, and an indeterminate bovid. Our material represents the first report of Eotragus from Southeast Asia, thus greatly extending the geographic distribution of this genus across the Old World continents. While comparisons of the Southeast Asian specimens with abundant material of E. clavatus from Sansan (France) and E. aff. clavatus from Tarazona de Aragón (Spain) indicate a high degree of intraspecific variation within single species of Eotragus, the existence of two distinct taxa at Mae Moh remains a possibility. Based on previous carbon isotope studies of Mae Moh herbivore tooth enamel, Eotragus lampangensis sp. nov. foraged predominantly in an ecotone between grassland and forest.
Although the suid genus Kolpochoerus is well known from the Plio-Pleistocene of Africa, the evolutionary history of one of its constituent species, K. majus, remained obscure until substantial fossil evidence accumulated during the last 20 years, largely from sites in Ethiopia. Here, we describe Kolpochoerus phillipi sp. nov., based on a fairly complete skull and the remains of additional individuals from ∼2.5 Ma deposits at Matabaietu, in the Middle Awash study area of Ethiopia. Based on a phylogenetic analysis, we suggest that K. phillipi sp. nov. belongs to a clade of “bunolophodont suines” including K. majus and the extant giant forest hog Hylochoerus meinertzhageni. Within this clade, K. phillipi sp. nov. likely represents a potential ancestor of K. majus, based on its morphology and stratigraphic position.
Extant porpoises (Phocoenidae) are odontocetes characterized by their small size, short and wide rostrum, late (or absent) completion of epiphyseal ankylosis in the vertebral column (= physical maturity), and short life cycles, all of which are thought to have resulted from progenetic evolution. We describe a small fossil phocoenid from the lower Pliocene Koetoi Formation of Hokkaido (northern Japan), preserving a small, narrow rostrum, as well as anteroposteriorly elongate thoracic and lumbar vertebral centra with completely fused epiphyses. Physical maturity in this specimen occurred significantly earlier than in extant phocoenids, as shown by dental data indicating that the specimen died at only four years of age. The difference between the present material and extant porpoises may be attributable to different growth rates during ontogeny. The long centra and caudally inclined neural spines of the specimen from Hokkaido are primitive characters among phocoenids. By contrast, the great height of its neural spines is highly derived, even among extant species, and suggestive of a fast swimmer. In terms of its vertebral morphology, the new specimen falls within a morphological continuum defined by the archaic Numataphocoena yamashitai and the highly derived vertebral morphology of Phocoenoides dalli. Phocoenid vertebral evolution has been complex and frequently convergent, as opposed to stepwise and unidirectional. The different vertebral morphologies of the new specimen and the contemporaneous extinct taxa Numataphocoena and Piscolithax longirostris indicate that they were adapted to different environments.
Oceanic dolphins (Odontoceti: Delphinidae) constitute the most speciose family of extant cetaceans, yet their fossil record is limited. Although several extinct species are known from Mediterranean and North Atlantic localities, there are few examples from deposits along the Pacific Rim. Despite the rich record of successive marine mammal fossil assemblages in the extensively sampled eastern North Pacific, only one fossil delphinid, Protoglobicephala (Pliocene, Baja California), has been described. We report globicephaline remains from the Mio-Pliocene Purisima Formation of Northern California, including a partial cranium and two isolated petrosals. The skull exhibits large ridges on the premaxillae, and cannot be referred to any extant globicephaline genus. Similarly, the petrosals cannot be referred to any described delphinid genus, although they are most similar to those of Globicephala. Linear regression analyses demonstrate that promontorium length and bony nares width scale significantly within delphinidans, and provide a new method for testing referrals of isolated fossil odontocete petrosals to taxa known only by crania. Applying this method to the new globicephalines from the Purisima Formation, we find the petrosals to be too small to represent the same taxon as the skull, thus indicating the presence of two separate species. Our results demonstrate that globicephalines had achieved a worldwide distribution by the early Pliocene, suggesting diversification of the subfamily by 5 Ma.
I describe Archaeonothos henkgodthelpi gen. et. sp. nov., a small (estimated body mass ∼40–80 g) tribosphenic metatherian from the early Eocene Tingamarra Fauna of southeastern Queensland, Australia. This taxon, known only from a single isolated upper molar (M2 or M3) is characterised by a very distinctive combination of dental features that, collectively, probably represent faunivorous adaptations. These include: a straight, elevated centrocrista; a metacone considerably taller than the paracone; a wide stylar shelf (∼50% of the total labiolingual width of the tooth); reduced stylar cusps; a long postmetacrista; a small and anteroposteriorly narrow protocone; an unbasined trigon; and the absence of conules. Some of these features are seen in dasyuromorphians, but detailed comparisons reveal key differences between A. henkgodthelpi and all known members of this clade. A. henkgodthelpi also predates recent molecular estimates for the divergence of crown-group Dasyuromorphia. Similar dental features are seen in a number of other metatherians, including the South American sparassodonts, Wirunodon chanku from the ?middle—late Eocene Santa Rosa local fauna of Peru, and Kasserinotherium tunisiense from the early Eocene Chambi fauna of Tunisia, although whether A. henkgodthelpi is closely related to any of these taxa is unclear based on available evidence. I therefore refer A. henkgodthelpi to Metatheria incertae sedis. Potential relatives of A. henkgodthelpi are unknown from any other Australian fossil deposit.
Isolated teeth from vertebrate microfossil localities often provide unique information on the biodiversity of ancient ecosystems that might otherwise remain unrecognized. Microfossil sampling is a particularly valuable tool for documenting taxa that are poorly represented in macrofossil surveys due to small body size, fragile skeletal structure, or relatively low ecosystem abundance. Because biodiversity patterns in the late Maastrichtian of North American are the primary data for a broad array of studies regarding non-avian dinosaur extinction in the terminal Cretaceous, intensive sampling on multiple scales is critical to understanding the nature of this event. We address theropod biodiversity in the Maastrichtian by examining teeth collected from the Hell Creek Formation locality that yielded FMNH PR 2081 (the Tyrannosaurus rex specimen “Sue”). Eight morphotypes (three previously undocumented) are identified in the sample, representing Tyrannosauridae, Dromaeosauridae, Troodontidae, and Avialae. Noticeably absent are teeth attributed to the morphotypes Richardoestesia and Paronychodon. Morphometric comparison to dromaeosaurid teeth from multiple Hell Creek and Lance formations microsites reveals two unique dromaeosaurid morphotypes bearing finer distal denticles than present on teeth of similar size, and also differences in crown shape in at least one of these. These findings suggest more dromaeosaurid taxa, and a higher Maastrichtian biodiversity, than previously appreciated.
A partial cranial endocast and right inner ear of the Cretaceous abelisaurid dinosaur Aucasaurus garridoi were digitally reconstructed from CT scans. The forebrain, midbrain, and hindbrain resemble the morphology described for the abelisaurids Majungasaurus and Indosaurus. However, Aucasaurus exhibits a floccular process that is relatively larger than that of Majungasaurus. In Aucasaurus the flocculus is enclosed in an 8-shaped floccular recess, similar in shape and size to that observed in Abelisaurus, suggesting that the two Patagonian taxa were capable of a slightly wider range of movements of the head. Here we describe the second inner ear known for the Abelisauridae. The labyrinth of the inner ear is similar in shape and size to the semicircular canals of Majungasaurus, although the lateral semicircular canal is shorter in Aucasaurus.
Many sauropod ghost lineages cross the Middle Jurassic, indicating a time interval that requires increased sampling. A wide taxonomic spectrum of sauropodomorphs is known from the Middle Jurassic of China, but the braincase of a new sauropod, named here Nebulasaurus taito gen. et sp. nov., is distinct. Nebulasaurus is sister taxon to Spinophorosaurus from the Middle Jurassic of Africa and represents a clade of basal eusauropods previously unknown from Asia. The revised faunal list indicates dramatic transitions in sauropodomorph faunas from the Jurassic to Cretaceous of Asia; these are consistent with geographic isolation of Asia through the Late Jurassic. Non-sauropod sauropodomorphs, non-mamenchisaurid eusauropods (including basal macronarians), and mamenchisaurids successively replaced previous grades through the Jurassic, and titanosauriforms excluded all other sauropod lineages across the Jurassic—Cretaceous boundary.
Previously unpublished trionychid turtle material from the Upper Cretaceous (Santonian—lower Campanian) Bostobe Formation from the Baybishe and Baykhozha localities in Kazakhstan is described. The material represents a new species of Khunnuchelys, a large, skull-based clade of Cretaceous Asian trionychids. Concordant with other partial skulls and fragmentary specimens described previously, Khunnuchelys lophorhothon sp. nov. has the unusual features of a beaklike maxilla and a vaulted, expanded triturating surface. In addition, the specimens reveal novel features including a constricted skull roof. Although estimates of the length of the carapace differ depending on estimation method, the skull belonged to a turtle of comparable size to the shell-based species “Trionyx” kansaiensis from the same formation. It is likely that K. lophorhothon and T. lophorhothonkansaiensis are synonymous, but this can be proved only by a find of associated skull and shell material.
Odontochelys semitestacea, the oldest known turtle, from the Late Triassic of China, shows a pathology. Sharply defined, focal depressions were noted on the articular surfaces of both humeri, documenting avascular necrosis. Diving habits of Mesozoic marine reptiles have been characterized on the basis of this localized form of bone death attributed to decompression syndrome. Pursuit by a predator was likely the cause of dangerously rapid depth changes by swimming turtles. The prevalence of avascular necrosis decreased geometrically from the Cretaceous to the Pleistocene. This study suggests that the habit of repetitive diving in turtles was already present in the Late Triassic, but that protective physiological and behavioral adaptations had not yet evolved.
Sphenacodontid synapsids were major components of early Permian ecosystems. Despite their abundance in the North American part of Pangaea, they are much rarer in Europe. Among the few described European taxa is Neosaurus cynodus, from the La Serre Horst, Eastern France. This species is represented by a single specimen, and its validity has been questioned. A detailed revision of its anatomy shows that sphenacodontids were also present in the Lodève Basin, Southern France. The presence of several synapomorphies of sphenacodontids—including the teardrop-shaped teeth—supports the assignment of the French material to the Sphenacodontidae, but it is too fragmentary for more precise identification. The discovery of sphenacodontids in the Viala Formation of the Lodève Basin provides additional information about their ecological preferences and environment, supporting the supposed semi-arid climate and floodplain setting of this formation. The Viala vertebrate assemblage includes aquatic branchiosaurs and xenacanthids, amphibious eryopoids, and terrestrial diadectids and sphenacodontids. This composition is very close to that of the contemporaneous assemblages of Texas and Oklahoma, once thought to be typical of North American lowland deposits, and thus supports the biogeographic affinities of North American and European continental early Permian ecosystems.
Whereas cosmopolitan distribution patterns are established for many Late Devonian vertebrates (e.g., placoderms, onychodontiforms), few palaeobiogeographic studies have considered chondrichthyans. Recent discoveries of shark material demonstrate that some chondrichthyans were cosmopolitan by the Middle Devonian. Abundant Givetian microremains have been recovered from the Cairo quarry in eastern New York State, USA. These include teeth of two shark species with Gondwanan affinities, the omalodontid Portalodus mannoliniae sp. nov. and the antarctilamnid Wellerodus priscus. Abundant teeth of P. mannoliniae sp. nov. are characterized by a smooth diplodont crown, polarized cusps, and a labially oriented base. The teeth demonstrate monognathic heterodonty. The juvenile morph is distinguished from the adult by smaller size, slender cusps, and variation in the shape of the base. W. priscus is represented by rare juvenile teeth. Two groups of scales that show affinity to material from northern (Spain) and East Gondwana (Antarctica) are tentatively attributed to the two described species. Antarctilamnid distribution suggests a north Gondwanan origin and a colonization of the margin of the landmass before dispersing to Laurentia by the Middle Devonian. This material further indicates that vertebrate global dispersal was initiated by the Middle Devonian, and emphasizes earlier palaeogeographic interpretations that the Middle Devonian “Hamilton fauna” of North American Laurussia originated in the Early Devonian in South American Gondwana.
A newly discovered trilobite fauna from the early to middle Permian Qarari Unit of northeastern Sultanate of Oman is described. It comprises exceptionally complete and well-preserved examples of five proetoid species, belonging to genera typical of an eastern Tethyan region extending through southern Asia to Timor. The shallow shelf fauna compares closely with one of Artinskian age from Afghanistan. Permian species previously assigned to Paladin are here considered unrelated to this predominantly Carboniferous clade, and are placed in a new ditomopygine genus, Simulopaladin, type species Simulopaladin tridentifer sp. nov. Three other ditomopygine species are described: Hentigia ornata sp. nov., Iranaspidion elephas sp. nov., and Acanthophillipsia felicitae sp. nov. The proetid Triproetus bonbon sp. nov. is the most complete material known of a more widespread genus.
The genus Laevitomaria is reviewed and its palaeobiogeographical history is reconstructed based on the re-examination of its type species L. problematica, the study of material stored at the National Natural History Museum of Luxembourg, and an extensive review of the literature. The systematic study allows ascribing to Laevitomaria a number of Jurassic species from the western European region formerly included in other pleurotomariid genera. The following new combinations are proposed: Laevitomaria allionta, L. amyntas, L. angulba, L. asurai, L. daityai, L. fasciata, L. gyroplata, L.isarensis, L. joannis, L. repeliniana, L. stoddarti, L. subplatyspira, and L.zonata. The genus, which was once considered as endemic of the central part of the western Tethys, shows an evolutionary and palaeogeographical history considerably more complex than previously assumed. It first appeared in the Late Sinemurian in the northern belt of the central western Tethys involved in the Neotethyan rifting, where it experienced a first radiation followed by an abrupt decline of diversity in the Toarcian. Species diversity increased again during Toarcian—Aalenian times in the southernmost part of western European shelf and a major radiation occurred during the Middle Aalenian to Early Bajocian in the northern Paris Basin and southern England. After a latest Bajocian collapse of diversity, Laevitomaria disappeared from both the central part of western Tethys and the European shelf. In the Bathonian, the genus appeared in the south-eastern margin of the Tethys where it lasted until the Oxfordian.
Early Jurassic aragonitic foraminifers are outstandingly well-preserved in the Marmorea crust, a multiphased ferromanganese layer limiting the Schnöll and Adnet formations (Adnet, Northern Calcareous Alps, Austria). This remarkable preservation, related to the pervasive impregnation of aragonitic tests prior to their recrystallization, allowed observing unknown diagnostic features of the genus Involutina, which typifies the Suborder Involutinina. Thanks to a detailed examination of the Adnet specimens, this paper clarifies the taxonomy, systematic position, and phylogeny of Involutina. A new diagnosis, structural model, and lineage are introduced for the group. Involutina is the direct descendant of Aulotortus and the two taxa probably showed a parallel evolution. As Aulotortus, Involutina presents a high intraspecific variability and its diversity must be revised downward. Current phylogenetic and taxonomic frames of the Suborder Involutinina are firmly questioned as, contrary to previous schemes, the type-genus possesses more than one lamellar deposit per whorl. In Involutina, the height and distribution of papillae on the test surface is not random and probably related to a biological function. We here propose that the papillose lamellae and tube infoldings that characterize representatives of the genus were rudimentary features for light catching and symbiont positioning, respectively.
The distribution of fossil wood genera has been demonstrated to be an effective proxy for Mesozoic terrestrial climates. In this study, we investigated the phytocoenoses, which were associated with Xenoxylon confirmed to be a marker for a cool and/or wet climate in a boreal hemisphere (i.e., Xenoxylon-phytocoenoses) during the Mesozoic, using specimens of fossil wood. It was confirmed that Xenoxylon co-occurs more often with some wood genera than with others. For example, Protocedroxylon, a wood that is most likely related to the Pinaceae, is the genus most often associated with Xenoxylon-phytocoenoses. Although Taxodioxylon is also found in Xenoxylon-phytocoenoses, it is not found, however, as consistently as Protocedroxylon. The distribution and diversity of Xenoxylon-phytocoenoses changed throughout the Mesozoic. During the Late Triassic and Late Cretaceous, Xenoxylon-phytocoenoses had low diversity and were restricted to higher palaeolatitudes during the Late Cretaceous. However, during the Early to Middle Jurassic, Xenoxylon-phytocoenoses were distributed much farther south, while their diversity concomitantly increased sharply. From the Late Jurassic to the Early Cretaceous, the distribution of Xenoxylon-phytocoenoses moved northward in Europe and even more so in East Asia. The changes in the distribution of Xenoxylon-phytocoenoses are in agreement with changes in both global and regional climates. Our results also demonstrated that, within the Xenoxylon distribution range, the corresponding phytocoenoses were differentiated along a latitudinal gradient and according to the global climate change patterns during the Mesozoic.
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