A recent study on the fossil history of North American woodrats and their relatives suggested that the Neotomini includes two subtribes: the Neotomina (woodrats; Tsaphanomys, Neotoma, Hodomys, Xenomys) and Galushamyina (reprats: Protorepomys, Galushamys, Miotomodon, Repomys, Nelsonia). The extinct Miocene Lindsaymys was proposed as a possible early neotominan, but not formally included in the Neotomini. In other studies, two extinct genera, Basirepomys and Paronychomys, occasionally have been treated as related to neotominans, but their ancestry had not been formally explored in detail. We performed a phylogenetic analysis on representatives of all genera with 40 dental and mandibular characters likely to be preserved in fossil material. The analysis resulted in a single most parsimonious tree supporting a neotominan-galushamyinan tribal classification of the Neotomini, and securely placed Lindsaymys within the Neotomina. None of the Basirepomys and Paronychomys species or Protorepomys bartlettensis nested within the Neotomini. The three Basirepomys species, possibly a paraphyletic group, are not closely related to Repomys, and Paronychomys (minus P. shotwelli = Tsaphanomys shotwelli) is not closely related to Onychomys, but is a possible sister group to the Neotomini. With the understanding that further study may remove it from Protorepomys, P. bartlettensis is tentatively retained within the genus, representing a relatively underived species possibly ancestral to both neotominan clades. Neotoma species of the extinct subgenus Paraneotoma and extant Neotoma may be paraphyletic at the generic level, but that determination will require further study.

The definition of the Cretaceous shark genus Cretodus Sokolov, 1965 is primarily based on isolated teeth. This genus includes five species. Among these, Cretodus houghtonorum Shimada and Everhart, 2019 is the only species based on a partially preserved skeleton. Here, the taxonomic attribution of a virtually complete skeleton of Cretodus from the Turonian of northeastern Italy is discussed, together with a few specimens from the Turonian of England. One of the latter is investigated through micropaleontological analysis to determine its stratigraphic position. The material is referred to Cretodus crassidens (Dixon, 1850), the diagnosis of which is emended herein. The dentition is tentatively reconstructed, exhibiting strong similarities with congeneric species, although it differs in having strong vertical folds on the main cusp labial face, a mesiodistally broad tooth aspect, weak and well-spaced ‘costulae’ at crown base, and a different dental formula in the number of parasymphyseal and lateral rows. Some tooth malformations are interpreted as feeding-related or senile characters. The Italian specimen suggests that Cretodus crassidens had a wide and laterally expanded mouth and head, a stout body, and attained a gigantic size. Cretodus crassidens was a moderate-speed swimming shark ecologically like the extant tiger shark Galeocerdo cuvier (Péron and Lesueur in Lesueur, 1822). The age estimate from vertebral-band counting suggests that the Italian individual was at least 23 years old and the growth model indicates a longevity of 64 years and a maximum attainable total length of 9–11 m. Cretodus crassidens occurs both in Boreal and Tethyan domains, implying a broad paleobiogeographic distribution and a preference toward offshore settings.

The fossil record of Early Triassic diapsids is very important to understand how the end-Permian mass extinction affected ecosystems and the patterns and processes involved in the subsequent biotic recovery. Vertebrate fossil assemblages of continental deposits in current-day South Africa, China, and Russia are the best source of information of this clade during the aftermath of the extinction event. Although considerably less sampled, the Induan continental rocks of the Panchet Formation of the Damodar Basin (eastern India) have also yielded a relatively diverse vertebrate assemblage composed of fishes, temnospondyls, synapsids, and a single proterosuchid taxon. Here, we report on a small isolated diapsid partial ilium (ISIR 1132) from the upper Panchet Formation. This specimen has a distinct morphology compared to other tetrapods that we know, including a shallow emargination on the dorsal margin of the anterior portion of the iliac blade, and ratio between height of iliac blade versus maximum height of iliac acetabulum at level of the dorsalmost extension of supraacetabular crest ≤0.45. Comparisons and a quantitative phylogenetic analysis found ISIR 1132 as a non-archosauromorph neodiapsid. This new specimen expands the reptile diversity in the Panchet Formation as well as for the rest of Gondwana, where Early Triassic non-archosauromorph neodiapsid species are extremely scarce.

Ohiocrinus byeongseoni n. sp. from the Middle Ordovician (Darriwilian) Jigunsan Formation of South Korea in the Sino-Korean (North China) block is the oldest species of Ohiocrinus of the Cincinnaticrinidae and the first record outside Laurentia. O. byeongseoni is characterized by a loosely clockwise-coiled anal sac, isotomous branching throughout arms, long, slender xenomorphic column, and small lichenocrinid-type holdfast. The new species occurs in association with a deep-water siliciclastic environment, unlike the Laurentian species with a shallow-water carbonate environment. The monospecific crinoid assemblage is interpreted as parautochthonous, considering that the crinoids were reworked by relatively weak down-current probably caused by storm but preserved within the environment where they lived. The occurrence of O. byeongseoni presents a considerable spatiotemporal gap and ecologic disparity in evolution of Ohiocrinus and the Cincinnaticrinidae.

UUID: http://zoobank.org/240417a4-8d33-4f45-aedb-afcc07f1ecbb

The Miocene beds of Kutch in India are well known for their mammalian assemblages, including the extinct ape Sivapithecus, but far less is known about the fossil squamates from this area. Although India with its over 800 reptile species is recognized as one of the global biodiversity hotspots, knowledge of past diversity and paleobiogeography of squamates on this subcontinent is very limited. We here report on new lizard finds, which have been recovered from two stratigraphic levels: the older Palasava locality (dated to the middle Miocene, ca. 14 Ma) and the younger Tapar site (late Miocene, ca. 11–10 Ma). Although fragmentarily preserved, the material described here sheds important light on the composition and paleobiogeography of squamates during the Miocene in South Asia. The older Palasava locality contains cf. Uromastyx s.l. and Varanus sp., the latter representing the oldest record of this taxon in the region of India south of the Himalayas and its occurrence here suggests a mean annual temperature not less than 15°C. The material from the younger Tapar locality consists of an unidentified acrodontan lizard, here questionably placed in agamids, and a skink. The latter shows a resemblance to mabuyines, however, the fragmentary nature of the material does not allow a precise allocation without doubts. The cosmopolitan mabuyines have been suggested to have their origin in Asia, so the potential presence of mabuyines in the Tapar locality might represent the first, but putative, Asian evidence of the occurrence of this group in the Miocene.

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.

A latest Paleocene charophyte flora collected from the South Gobi area in the Junggar Basin, western China, includes the geographically widespread taxa Peckichara torulosa var. varians (Dollfus and Fritel, 1919) Sanjuan, Vicente, and Eaton, 2020, Lychnothmanus vectensis (Groves, 1926) Soulié-Märsche, 1989, and Gyrogona lemani capitata Grambast and Grambast-Fessard, 1981. Lychnothmanus vectensis (as Lychnothmanus aff. L. vectensis) is known from the Cretaceous–Paleocene transition in eastern China and the latest Paleocene in western China, with likely additional records from the United States (Utah). The earliest European records of L. vectensis are from the late Eocene to early Oligocene in Spain, France, and England. Similarly, the oldest record of G. lemani capitata is from the latest Paleocene in the South Gobi area, with younger records from the middle Eocene of France. These latest Paleocene gyrogonite assemblages demonstrate the origin of these charophyte lineages in Asia. The dispersal of these charophytes from Asia to Europe in the middle to late Eocene appears to have occurred before the retreat of the Turgai Strait in both the Tarim area and the Siberian Basin by the end of the late Eocene and before the “Grande Coupure” in Europe and the Mongolian Remodelling in Asia during the Eocene–Oligocene transition. We hypothesize that waterbirds may have facilitated this intercontinental dispersal, and that idea is supported by the shared occurrence of avian groups in Central Asia and Europe in the middle and late Eocene.

Carnivores make traces on bones with their teeth when feeding. A true predatory bite trace (predichnia) forms when a predator catches and kills its prey or attempts to do so. Both predators and scavengers may leave their nonpredatory feeding traces during postmortem food processing. Despite the interpretative uncertainties as to the ethology such ichnofossils may represent, the bite traces have been traditionally classified as predichnia—traces of predation. Previously, there was no alternative ethological category available for workers to classify them. The present paper fills that gap and describes tooth-made ichnofossils from the continental Upper Triassic Grabowa Formation of southern Poland. It discusses modes the serration and striations might have formed along Linichnus edges, potential makers of the trace fossils, feeding strategies, and food-processing behaviors the ichnites may represent. All the bite traces are thought to act as a record of carnivorous behaviors and are classified as sarcophagichnia, a new ethological category (traces of feeding on a body). Finally, all the studied bite traces were likely inflicted postmortem and are classified as necrophagichnia (traces of feeding on an already dead body), most likely produced by scavengers in the studied cases. Data on recent carnivores link these ichnites with postmortem food-processing behaviors, such as dismembering and defleshing. Scavenging could, in fact, have been a preferred carnivorous feeding strategy in the seasonal Norian climate of the area. Dry seasons could have perhaps increased vertebrate mortality rates and provided plenty of carcasses for carnivores to feed on.

We report new specimens of the Plotopteridae from Washington State (USA), an area where these flightless seabirds underwent significant diversification during the late Eocene and Oligocene. To date, five plotopterid species from western Washington have been formally named. Specimens previously assigned to Tonsala buchanani Dyke, Wang, and Habib, 2011 belong to at least two, but probably even three, different species. One of these, the large-sized “Whiskey Creek specimen” from late Eocene deposits mapped as the Makah Formation, is the oldest known plotopterid and is here tentatively assigned to ?Klallamornis clarki Mayr and Goedert, 2016. Another specimen originally referred to T. buchanani is also likely to belong to a different species and is among the most substantial records for North American plotopterids. We formally transfer T. buchanani to the taxon Klallamornis and show that the only unambiguously identified specimen of the species—the holotype—is currently poorly diagnosed from Klallamornis abyssa Mayr and Goedert, 2016, which is from coeval strata of the Pysht Formation. Although the holotype of K. abyssa is larger than that of K. buchanani, there remains a possibility that plotopterids were sexually dimorphic in size. We describe the first ungual phalanx of a plotopterid, which is referred to K. buchanani, and report previously unknown elements of the large ?K. clarki and the first records of this species from the Lincoln Creek Formation. Current data indicate that plotopterids originated in the middle or late Eocene on islands off western North America, and we hypothesize that the radiation of these birds in the North Pacific Basin may have been related to the evolution of kelp forests.

A new specimen of Basilosaurus cetoides was discovered on the banks of the Flint River in Albany, Georgia, USA, in 2010. This fossil, which was the most complete specimen of the species from Georgia to date, consisted of five nearly complete and two partial post-thoracic vertebrae, tentatively identified as S4 through Ca6. During excavation, however, the site was looted and most of the specimen was lost to science. Nonetheless, we use this discovery as an opportunity to update the current state of knowledge on the stratigraphic, biogeographic, and environmental distribution of Basilosaurus in North America, as well as the position of the late Eocene shoreline in the southeastern United States. The results show that Basilosaurus was most abundant across the southeastern coastal plain during the early to middle Priabonian, coincident with the late Eocene maximum marine transgression. The decline in Basilosaurus localities is associated with the retreating shoreline of the terminal Eocene. The majority of Basilosaurus localities fall well south of the position of the late Eocene shoreline hypothesized in this study, suggesting the genus favored middle to outer neritic zones of the epicontinental sea. The comparatively low number of Priabonian specimens in the Atlantic Coastal Plain versus the Gulf Coastal Plain, then, suggests the presence of shallow zones in the Atlantic Coastal Plain that may have limited the distribution of Basilosaurus across the region. The hypothesized shoreline of this study ultimately differs from earlier reconstructions by extending the Mississippi embayment at the Bartonian/Priabonian boundary farther north than previously noted.

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.

UUID: http://www.zoobank.org/25d9b772-bd7d-4ad6-bfc6-ba02b1567cf3

Modular coral-like fossils occur in thrombolitic reefal beds at two stratigraphic levels within the Lower Ordovician (Floian) Barbace Cove Member of the Boat Harbour Formation, in the St. George Group of western Newfoundland. They are here assigned to Reptamsassia n. gen.; R. divergens n. gen. n. sp. is present at both levels, whereas a comparatively small-module species, R. minuta n. gen. n. sp., is confined to the upper level. Reptamsassia n. gen. resembles the Ordovician genus Amsassia in its phacelocerioid structure, back-to-back walls of adjoining modules, module increase by longitudinal fission involving infoldings of the wall, tabula-like structures that are continuous with the vertical module wall, and calices with concave-up bottoms. The new genus is differentiated by its encrusting habit, modules with highly variable growth directions and shapes throughout skeletal growth, and modules that may separate slightly or diverge from one another following fission. Together, Amsassia and Reptamsassia n. gen. are considered to represent a distinct group of calcareous algae, the Amsassiaceae n. fam., which possibly belongs to the green algae. The Early Ordovician origination of Amsassia followed by Reptamsassia n. gen. contributed to the beginning of the rise in diversity on a global scale and in reefal settings during the Great Ordovician Biodiversification Event. Reptamsassia minuta n. gen. n. sp. was an obligate symbiont that colonized living areas on its host, R. divergens n. gen. n. sp., with isolated modules of R. divergens n. gen. n. sp. able to persist in the resulting intergrowth with R. minuta n. gen. n. sp. This is the earliest known symbiotic intergrowth of macroscopic modular species, exemplifying the development of ecologic specialization and ecosystem complexity in Early Ordovician reefs.

Parioxys ferricolus Cope, 1878 is a long-neglected taxon of dissorophid temnospondyls from the early Permian (Cisuralian) of Texas. Reexamination of the original material and preparation of a previously undescribed specimen shed some light on the ontogeny and reveal numerous eucacopine features. P. ferricolus is characterized by: (1) a preorbital region 1.5 times as long as the posterior skull table in juveniles and twice as long in adults, (2) a parietal only two-thirds the length of the frontal, (3) interpterygoid vacuities shorter than half the skull length, (4) choanae posteriorly extended, and (5) vomer with enlarged, V-shaped median depression. P. ferricolus is found nested with Kamacops acervalis Gubin, 1980, with which it shares synapomorphies in the palate, within the dissorophid clade Eucacopinae.

With differentiated tissues and organs, a high-level eukaryotic macroalga Lanceaphyton xiaojiangensis n. gen. n. sp. lived on the middle–late Ediacaran (ca. 560–551 Ma) seafloor in South China. Its body had a pith (perhaps mechanical tissue) and outer tissue (perhaps epidermis and/or cortex). The lance-like macroalga consists of an unbranching thallus that grew over the sediment surface for sunlight and a holdfast grown into sediments to keep the thallus fixed on the seafloor. The pithy stipe (lower thallus) might have served to support the upper pithless thallus for photosynthesis. The holdfast is composed of a tapering pithy rhizome growing down into the sediments, with many filamentous pithless rhizoids dispersedly growing within the sediments. With the differentiated tissues and organs, especially the pith accounting for about half of the width of the rhizome and stipe, Lanceaphyton n. gen. was a high-level eukaryotic macroalga, similar to phaeophytes in morphological features, but further research is needed on its microstructural details. The pithy macroalga shows that the macroalgal pith had emerged in the Ediacaran.

UUID: http://zoobank.org/bc924c5c-84e4-4170-9ca1-caee0d56c6d5.

UUID: http://zoobank.org/bae1f4cb-c5c1-4dfd-85e2-22bfc929e4a0

One of the oldest fossil horseshoe crabs figured in the literature is Entomolithus lunatus Martin, 1809, a Carboniferous species included in his Petrificata Derbiensia. While the species has generally been included within the genus Belinurus Bronn, 1839, it was recently used as the type species of the new genus Parabelinurus Lamsdell, 2020. However, recent investigation as to the appropriate authority for Belinurus (see Lamsdell and Clapham, 2021) revealed that all the names in Petrificata Derbiensia were suppressed in Opinion 231 of the International Commission on Zoological Nomenclature (1954) for being consistently nonbinomial under Article 11.4 of the International Code of Zoological Nomenclature (ICZN) (International Commission on Zoological Nomenclature, 1999). Despite the validation of several species names for anthozoans, brachiopods, and cephalopods described in Petrificata Derbiensia in subsequent rulings (International Commission on Zoological Nomenclature, 1956a, b), Belinurus lunatus has not been the subject of any subsequent Commission ruling or opinion, and so its use in Petrificata Derbiensia remains suppressed. The Belinurus lunatus species name was used in several subsequent publications during the 1800s, none of which made the name available under ICZN article 11.5; Parkinson (1811) is also suppressed for being nonbinomial, while Woodward (1830), Buckland (1837), Bronn (1839), and Baily (1859) refer to the species only as a synonym of Belinurus trilobitoides (Buckland, 1837) through citation to the suppressed Pretificata Derbiensia. The first author to make Belinurus lunatus an available name was Baldwin (1905), who used the name in reference to a new figured specimen from Sparth Bottoms, Rochdale, UK, but again as an explicit junior synonym of Belinurus trilobitoides (Buckland, 1837). Therefore, it was not until Eller (1938) treated B. lunatus as a distinct species from B. trilobitoides that B. lunatus became an available name as per ICZN Article 11.6.1 under the authorship of Baldwin (1905) following ICZN Article 50.7.

Studies of molecular biology suggest a deep origin of coralline red algae in the Neoproterozoic, but unequivocal representatives have not been found in Precambrian rocks. Such scarcity in the fossil record significantly limits our knowledge regarding the origin and early evolution of this group. Phosphatized, fragmented columnar microfossils from the Zhenba microfossil assemblage at the base of the Ediacaran Dengying Formation, South China are presented. Morphologically they represent a novel taxon characterized by: (1) variable and irregularly shaped cross sections, and (2) discontinuous, nonisopachous laminae transversely stacked inside the columns. Integrated morphological comparisons favor their interpretation as probable erect coralline algae. Anatomically, the Zhenba columns resemble present-day geniculate coralline algae in columnar thalli and stromatolithic tissue construction, although they differ from modern geniculate corallines in lacking highly calcified skeletons and tissue-organ differentiation. These anatomical similarities collectively suggest that the Zhenba columns probably represent stem-group coralline algae. The new finding indicates that early coralline algae could be more diverse than previously thought, and their initial morphological diversification and resultant ecological expansion probably occurred in the middle to late Ediacaran.

Correct identification of fossil taxa is immensely important for dating molecular phylogenies and understanding when and how quickly modern biodiversity evolved. Fossils that are available for a clade of interest and can be directly incorporated in the phylogenetic analysis are considered primary sources of time calibration, whereas calibrations inferred from other studies are secondary (Arroyave et al., 2013). Studies of taxonomic groups that lack fossils must either expand their analyses to include fossilized outgroup lineages, use secondary calibrations, or use more problematic primary calibrations, e.g., vicariant geologic events. The use of vicariant geologic events to calibrate phylogenies poses the risk of circular reasoning, because the goal of many such studies is to determine how geologic events have affected diversification. Near et al. (2012) argued that fossil calibrations external to clades of interest, but still within the broader Actinopterygian (ray-finned fishes) tree, could be used as means of calibrating a generalized molecular clock, but internal calibrations are still valuable for refining such inferences (Arroyave et al., 2013).

A new enantiornithine, Musivavis amabilis n. gen. n. sp., is reported from the Lower Cretaceous Jehol Biota in western Liaoning, China. The new taxon is similar to the bohaiornithids in the robust subconical teeth, bluntly expanded omal ends of the furcula, caudolaterally oriented lateral trabeculae with triangular distal ends of the sternum, and a robust second pedal digit. Yet it differs from members of Bohaiornithidae in several features recalling other enantiornithine lineages, such as the acuminate rostral ramus of maxilla, the shape of the coracoid lateral margin, the presence of craniolateral processes on the sternum, the proportions of the manual phalanges, and the unspecialized third pedal ungual phalanx. A comprehensive phylogenetic analysis of Mesozoic birds shows that homoplasy significantly affects the reconstruction of enantiornithine relationships. When all phylogenetic characters are considered of equal weight, Musivavis is reconstructed in a lineage related to a radiation of large-bodied enantiornithines including Bohaiornithidae and Pengornithidae. Alternative scenarios based on progressive downweighting of the homoplastic characters support more basal placements of the pengornithids among Enantiornithes, but do not alter the affinity of Musivavis as a member of the “bohaiornithid-grade” group.

UUID: http://zoobank.org/617c7062-21ab-4d33-ae80-4edf5a129683