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We describe here the early Spathian (Early Triassic) Paris Biota decapod fauna from the western USA basin. This fauna contains two taxa of Aegeridae (Dendobranchiata), namely Anisaeger longirostrus n. sp. and Aeger sp. that are the oldest known representatives of their family, thus extending its temporal range by 5 Myr back into the Early Triassic. This fauna also includes two representatives of Glypheida (Pleocyemata) with Litogaster turnbullensis and Pemphix krumenackeri n. sp., confirming for the former and extending for the latter the temporal ranges of their respective superfamilies back to the Early Triassic. Overall, the Paris Biota decapods are some of the oldest known representatives of Decapoda, filling in an important gap in the evolutionary history of this group, especially during the Triassic that marks the early diversification of this clade. Additionally, we compile and provide overviews for all known Triassic decapods, which leads to the revision of four species of Middle and Late Triassic Aegeridae, and to a revised family assignment of a Middle Triassic Glypheida. Based on this refined dataset, we also investigate decapod diversity throughout the Triassic. We show that the apparent increase in decapod taxonomic richness is probably driven by the heterogeneity of the fossil record and/or sampling effort, and that the decapod alpha diversity is actually relatively high as soon as the Early Triassic and remains rather stable throughout the Triassic.
The Cambrian successions at the Chihuarruita Hill outcrop, Sonora, Mexico, have yielded two successive linguliform brachiopod assemblages that are transitional between Cambrian Stage 4 and the newly recognized global Wuliuan Stage. The lowermost assemblage includes Dictyonina sp., Paterina sp., Eothele sp., Hadrotreta rara? (Cooper), and Linnarssonia arellanoi? (Cooper), coming from the upper part of the Buelna Formation. The younger, recently named El Gavilán Formation contains a more diverse linguliform brachiopod assemblage, including Acrothele concava Cooper, Batenevotreta? mexicana n. sp., Dictyonina minutipuncta Cooper, Eothele sp., Eoobolus sp., Hadrotreta rara? (Cooper), Linnarssonia arellanoi? (Cooper), Micromitra sp., Paterina sp., and Prototreta sp. The El Gavilán Formation contains a diverse trilobite fauna suggesting Delamaran age in terms of the Laurentian regional stratigraphical scheme. The base of the global Wuliuan Stage and Miaolingian Series is defined by the first occurrence of Oryctocephalus indicus; in the absence of the index species, the base should be provisionally placed at the base of the El Gavilán Formation. The Wuliuan age of the brachiopod assemblage recovered from the El Gavilán Formation is supported by the occurrence of Acrothele in the Cambrian biostratigraphical succession of Himalaya, where the genus makes its first appearance in the Kaotaia prachina Zone. In addition, the co-occurrences of Acrothele and Eothele can be taken as an indication of the Wuliuan age of the fauna. A new biogeographic analysis confirms that the Eothele Fauna first appeared at the end of Cambrian Stage 4, as a result of increased faunal migration within the southern tropical latitudes directed from Australasian Gondwana to Laurentia.
Clathrodictyids are the most abundant stromatoporoids in the Upper Ordovician Xiazhen Formation (middle to upper Katian) of South China. A total of nine species belonging to four clathrodictyid genera are identified in the formation, including Clathrodictyon idense Webby and Banks, 1976, Clathrodictyon cf. Cl. microundulatum Nestor, 1964, Clathrodictyon cf. Cl. mammillatum (Schmidt, 1858), Clathrodictyon megalamellatum Jeon n. sp., Clathrodictyon plicatum Webby and Banks, 1976, Ecclimadictyon nestori Webby, 1969, Ecclimadictyon undatum Webby and Banks, 1976, Camptodictyon amzassensis (Khalfina, 1960), and Labyrinthodictyon cascum (Webby and Morris, 1976). The clathrodictyid fauna in the Xiazhen Formation is very similar to those from both New South Wales and Tasmania, although the latter two Australian regions do not share any common clathrodictyid species during the Late Ordovician. The paleobiogeographic pattern indicates that the northward drift of South China resulted in a favorable environment for the migration of clathrodictyids from other peri-Gondwanan terranes to South China. In addition, these peri-Gondwanan clathrodictyid species hosted various endobionts, representing a variety of paleoecological interactions. The high abundance and species-level diversity of clathrodictyid species presumably increased the substrate availability of suitable host taxa, judging from the diverse intergrowth associations between clathrodictyids and other benthic organisms. These paleoecological interactions between stromatoporoid and other organisms are known from the Late Ordovician and became more abundant and widespread in the Siluro–Devonian. Overall, the Late Ordovician clathrodictyid assemblage in South China demonstrates one of the highest species-level diversities among all peri-Gondwanan terranes and represents a precursor of the complex, clathrodictyid-dominated communities of later metazoan reefs during the Great Ordovician Biodiversification Event.
The fossil record of treeshrews, hedgehogs, and other micromammals from the Lower Siwaliks of India is sparse. Here, we report on a new genus and species of fossil treeshrew, specimens of the hedgehog Galerix, and other micromammals from the middle Miocene (Lower Siwalik) deposits surrounding Ramnagar (Udhampur District, Jammu and Kashmir), at a fossil locality known as Dehari. The treeshrew from Dehari (Sivatupaia ramnagarensis n. gen. n. sp.) currently represents the oldest record of fossil tupaiids in the Siwaliks, extending their time range by ca. 2.5–4.0 Myr in the region. Dietary analyses suggest that the new tupaiid was likely adapted for a less mechanically challenging or more frugivorous diet compared to other extant and fossil tupaiids. The occurrence of Galerix has only been recently documented from the Indian Siwaliks and the Dehari specimens help establish the likely presence of a relatively large Siwalik Galerix species in the Ramnagar region. In addition to the new treeshrew and hedgehogs, new specimens of the rodents Kanisamys indicus, Sayimys sivalensis, and Murinae indet. from Dehari help confirm that age estimates for the Ramnagar region are equivalent to the Chinji Formation in Pakistan, most likely corresponding to the middle to upper part of the Chinji Formation.
The diversity of mid-Cretaceous tyrannosauroids is poorly understood. We describe a partial tyrannosauroid femur from the Albian–Cenomanian Wayan Formation of eastern Idaho that helps to fill in an important spatiotemporal gap in the North American record of tyrannosaurs. This specimen, consisting of the proximal half of the bone, is morphologically similar to the femur of Moros intrepidus, a small-bodied tyrannosauroid from the Cenomanian Mussentuchit Member of the Cedar Mountain Formation of Utah, but not referable to this taxon. The Wayan femur lacks an autapomorphy diagnostic for Moros intrepidus, indicating the presence of a previously unrecognized tyrannosauroid taxon in the early Late Cretaceous of Laramidia. Histological results indicate that, at the time of death, this individual was at least five years old, skeletally immature, and undergoing growth at a moderate rate. The addition of this tyrannosauroid to the Wayan-Vaughn Assemblage provides additional evidence for the widespread distribution of various tyrannosauroid taxa in Laramidia during the early Late Cretaceous.
Theridiidae is a family of spiders with the fourth highest richness of extant species worldwide. Fossil Theridiidae is also the family of spiders with the largest number of Cretaceous, Eocene, and Miocene representatives. Phycosoma Pickard-Cambridge, 1880 is a theridiid genus without a fossil representative. In this study, we describe a new fossil species of Phycosoma, P. icti† n. sp., from an adult male specimen found in a piece of amber from the lower Miocene (ca. 23 Ma), from the Montecristo mines near Simojovel de Allende, Chiapas, Mexico. Phycosoma icti† n. sp. can be distinguished from all other Phycosoma males except P. corrugum by the long embolus partly covered by the conductor. In addition, it can be distinguished from P. corrugum by the oblique and constricted median apophysis; the narrow and pointed conductor; the long, curved, and bent theridiid tegular apophysis in its distal portion; and the prosoma ringed in the middle and along the ventral margin. Phycosoma icti† n. sp. represents the oldest recovered record of the genus. Extant Phycosoma species are still present in Chiapas, suggesting an extended evolutionary history of the genus in the region despite major geological changes over this time span in North America.
In this contribution, fossil woods from the Valle de La Luna Member of the Upper Triassic Ischigualasto Formation at Ischigualasto Provincial Park, San Juan Province, Argentina, are described. The specimens are preserved as silica permineralization in tuffs intercalated with carbonaceous mudstone beds interpreted as distal floodplain facies. The fossil woods were assigned to the new species Agathoxylon argentinum since their anatomy differs from the known Mesozoic Gondwanan species of the genus Agathoxylon. The combination of characters present in the new taxon indicates an affiliation with the conifer family Araucariaceae. Signals of fungal-mediated wood decay were observed, comparable to the activity of basidiomycetes. Spherical structures attached to the walls of the tracheids were recognized and are interpreted as holocarpic chytrid fungi. The growth rings were quantitatively analyzed. Low values of percentage diminution, percentage latewood, and Ring Markedness Index, and a mean percentage skew of +11.5, were obtained, suggesting that the new species was an evergreen gymnosperm. The stratigraphic distribution and taxonomic composition of the Ischigualasto Formation fossil-plant-bearing levels were studied. A vegetation change is recorded in the fossil level bearing Agathoxylon argentinum n. sp., marked by the replacement of the corystosperm genera and a diminution of arboreal corystosperms. This floristic change, in addition to other evidence, indicates humid paleoclimatic conditions for the uppermost part of the Valle de La Luna Member of the Ischigualasto Formation.
Borophagus is the terminal genus of the highly diverse and successful subfamily, Borophaginae. Skeletal remains of this bone-crushing canid are most commonly found in transitional or grassland environments across North America between Late Miocene–Middle Pleistocene, but are rare or absent in forested habitats. Here, we describe a humerus from the Gray Fossil Site of Tennessee, which is the first occurrence of this genus in a heavily forested ecosystem. The distinct limb proportions of Borophagus suggest the genus may have been well suited for a closed habitat like the Gray Fossil Site, contrary to where a majority of their fossils have been previously found. This discovery documents the first pre-Pleistocene occurrence of a canid in the Appalachian region of the eastern United States.
The Suwannee Limestone of Florida and Georgia, USA, is a carbonate paleoenvironment with a rich diversity of marine fossils. These record the presence of coral- and seagrass-bearing communities in the western Atlantic from the Eocene/Oligocene boundary to the mid Oligocene (33.9–28.1 Ma). From the Suwannee Limestone, we describe two new species of the cosmopolitan, diverse, and abundant coral genus Acropora (“staghorn corals”), Acropora suwanneensis new species and Acropora upchurchi new species, which predate all other described Acropora species from the Americas and Caribbean by at least 10 million years. Diminutive skeletal structure in both species is indicative of their living in protected calm-water habitats. At their time of occurrence, an Atlantic–Pacific connection was open through Central America. Both species belong to morphological lineages also present in the fossil record of the eastern Atlantic (Europe, or the western Tethys). These lineages no longer occur in the Atlantic Ocean, but they are extant in the Indo-Pacific region, with the closest modern relative of each new species occurring in the Pacific Ocean to central Indo-Pacific. Some species in the two morphological lineages have existed for up to 20+ million years. The new species may have played a part in perpetuating a cosmopolitan Tethyan fauna and expanding Acropora habitats in the western Atlantic, by which they likely also contributed to the renowned species and habitat diversity of Acropora in the modern Indo-Pacific.
Sebecidae is a clade of large carnivorous crocodyliforms that thrived in the Cenozoic and is the only lineage of the diverse and terrestrial group Notosuchia that survived the end-Cretaceous mass extinction event. Sebecus icaeorhinus is the best-known taxon from this clade, both in terms of its cranial and postcranial anatomy (known primarily from the holotype and specimen MPEF-PV 1776, respectively). Additional material represented by a partial skull (MMP 235) is the only specimen that has preserved a complete choanal region. We describe new information from this specimen based on an X-ray computed tomography and identify through comparisons with other taxa a large degree of variability in the palatal anatomy within Sebecidae, in particular in the shape and extension of the palatine (the bone that defines the anterior position and shape of the secondary choana). We quantify variation in the shape of the palatine bone of sebecids through a 2D morphogeometric analysis within the context of notosuchian crocodyliforms. Although traditional accounts of palatal evolution in crocodyliforms linked variation of this structure to the adaptation to the aquatic environment, our analysis allows recognition of eight palatine morphotypes among terrestrial crocodyliforms with very distinct paleoecological traits, including carnivorous, omnivorous, and possibly herbivorous taxa. Furthermore, we show that sebecids had a higher morphological disparity in the choanal region than other terrestrial groups of Notosuchia, underscoring the importance of this region for comparative, morphofunctional, and phylogenetic studies.
Antiarch placoderm fishes were an abundant component of the Middle Paleozoic vertebrate assemblages. Despite a large number of known taxa and specimens, the morphology and function of the skeletal elements of their jaws is inadequately known. Because of this, questions regarding their feeding modes and their roles in the trophic webs remain open. We present a skeletomuscular model of the antiarch jaw apparatus with an attempt to reconstruct its potential biomechanical function. The position of the upper jaw suborbital bones within the plane of the ventral side of the fish armor is suggested to represent the natural “mouth closed” position. During mouth opening, the suborbitals rotated rostrally with simultaneous depression and inward rotation of the infragnathals. The ball-and-socket jaw articulation might ensure this combined movement. Recently described lower jaw elements of Livnolepis zadonica (Obrucheva, 1983) and Bothriolepis sp. from the Upper Devonian (lower Famennian) of Central Russia demonstrating very deep and porous blades of the oral division of the infragnathals queried the structure of these bones in other antiarchs. Observed porosity reflects intense vascularization to supply blood to a connective tissue underlying a supposed keratinous sheath, which protected and strengthened the jaws, as well as made possible scraping tough food objects, such as thallus algae, from the substrate.
Having evolved during the Silurian in the Pan-Cathaysian zoogeographical province, antiarchs migrated to Gondwana during the Emsian and later to Euramerica during the Eifelian. Supposedly, antiarchs became the first macrophytophagous vertebrates occupying the trophic level of primary consumers during the late Silurian–Early Devonian. This event diversified the only previously existing predator–prey interrelationships between filter-feeding agnathans and predatory gnathostomes.
The trilobite Liostracina has been recognized as important in taxonomic and stratigraphic studies for more than a century. Until now, the genus Liostracina and family Liostracinidae have been known from only incomplete holaspid material, a degree 2 meraspis, and protaspides. A new locality in the Longha Formation (Cambrian: Guzhangian) of southeastern Yunnan, China, yields a rich collection of articulated holaspid exoskeletons and disarticulated sclerites of a new species, Liostracina fuluensis n. sp. These specimens reveal thoracic and ventral morphology that was previously unknown for Liostracina; they demonstrate that it has a rostral plate plus rostral and connective sutures, rather than a ventral median suture on the cephalic doublure. This confirms a natant hypostomal condition for the genus. Holaspid exoskeletal features, combined with evidence of a non-asaphoid-type protaspis, indicate that Liostracina is neither a primitive trinucleoid nor a primitive asaphoid. The family Liostracinidae is excluded from the superfamily Trinucleoidea and the order Trinucleida and reassigned to the order Ptychopariida.
The small, immature brachyopid stereospondyl Platycepsion wilkinsoni from the Early or Middle Triassic of Gosford, New South Wales, is redescribed from a developmental point of view. Whereas the sutural contact between postorbital and parietal suggested by earlier authors cannot be confirmed, a new autapomorphy, the posterior process of the interclavicle with a slightly concave posterior margin, is recognized. A further apomorphic state may be the probable presence of five pairs of ossified ceratobranchials. Soft-tissue preservation of external gills demonstrates that Platycepsion wilkinsoni represents a true larva, forming the first evidence of a larval stage in stereospondyls. Morphology and developmental stage of Platycepsion wilkinsoni indicates that brachyopids conserved the plesiomorphic ontogenetic pattern seen in most Paleozoic temnospondyls, including the early ossification of the dermal skull roof and delayed ossification of the postcranium. The strongly developed ornament, the widened lateral lines, and the well-ossified exoccipitals and stapes suggest an even faster ossification and differentiation of the skull compared to Paleozoic temnospondyls and might represent a general stereospondyl characteristic. In the vertebral column, only the poorly differentiated, still separate neural arches are ossified, whereas the centra remained cartilaginous. This implies that the plesiomorphic pattern of vertebral development was conserved in Mesozoic stereospondyls, irrespective of whether the adult vertebral morphology was rhachitomous, stereospondylous, or diplospondylous. The low degree of ossification of the postcranium, its slow differentiation, and the delayed development of the scapulocoracoid indicate an aquatic mode of life, and the strongly ossified skull and hyobranchial apparatus suggest an emphasis on powerful suction feeding.
The Late Ordovician (late Katian) Tcherskidium fauna consisted of large- and thick-shelled virgianid pentamerid brachiopods characterized by large and ribbed shells of Tcherskidium and Proconchidium and usually associated with Holorhynchus, Deloprosopus, and Eoconchidium. This unique fauna was widely distributed across several tectonic plates, largely confined to the paleoequatorial and especially the northern paleotropical zones, such as northern Laurentia, accretionary terranes of Alaska, Kolyma, Baltica, Siberia, Kazakh and adjacent terranes, and South China. In Laurentia, the eponymous genus Tcherskidium was predominant in regions north of the paleoequator and, in sharp contrast, was absent south of the paleoequator. In this study, Tcherskidium lonei n. sp. and Proconchidium schleyi n. sp. are described from Alaska and North Greenland, respectively, adding new data on the Tcherskidium fauna of the Late Ordovician Northern Hemisphere. Shell gigantism, together with the sharp paleobiogeographic division, suggests that the Late Ordovician (late Katian) Northern Hemisphere had a prevailing warm-water mass, probably due to the lack of large landmass beyond the northern tropics. This was in sharp contrast to the Southern Hemisphere, which was frequently influenced by cold-water invasions from the ice-bearing Gondwana supercontinent centered on the South Pole.