Members of the echinoid order Spatangoida, a highly diverse and abundant marine invertebrate clade, were important denizens of the Cretaceous Western Interior Seaway (WIS), an epicontinental seaway that divided North America in two during an interval of greenhouse conditions between roughly 100 and 65 million years ago. A phylogenetic analysis of spatangoids was conducted using a character matrix of 32 characters from 21 species. Species that occur in the WIS were considered comprehensively, and species from other regions such as South America, Europe, and North Africa were also incorporated into the analysis. Phylogenetic patterns retrieved are largely congruent with preexisting family-level classifications; however, species within several genera, especially Hemiaster and Heteraster, need to be reassigned so that classification better reflects phylogeny. The genera Washitaster and Heteraster are closely related, as are Mecaster, Palhemiaster, and Proraster; Pliotoxaster, Macraster, and Hemiaster; and Micraster and Diplodetus. Biogeographic patterns were also considered using the phylogeny, and several episodes of vicariance and range expansion were identified. These were possibly related to some of the various major episodes of sea-level rise and fall during the Cretaceous. In particular, Valangian–mid-Aptian regressions may have caused vicariance within Heteraster and Washitaster while other early spatangoid vicariance may be related to regressions during the late Aptian–early Cenomanian. Further, vicariance caused by regressions during the mid-Cenomanian–Maastrichtian may have driven diversification within Micraster and Diplodetus. Last, transgressions during the late Aptian–early Cenomanian seem to have spurred prominent range expansions in Mecaster and Hemiaster.

Ichthyerpeton bradleyae (Huxley in Wright and Huxley, 1866) is one of the seven tetrapods originally described by Huxley from the Jarrow Assemblage (Pennsylvanian, Langsettian Regional Substage equated with the Bashkirian International Stage) in south-eastern Ireland. The holotype, one of only two specimens considered to represent the taxon, consists of the postcranial skeleton, which has been highly compressed and has undergone extensive replacement of bone by carbonaceous material. The holotype is studied using microcomputed tomography, which reveals that the vertebral column has at least 25 diplospondylous vertebrae with cylindrical centra. Neural arches and a haemal arch are described for the first time. Neural arches in the caudal region are paired and neural spines only contact one another dorsally. The hemal arch is fused and wraps around the ventral margin of the centrum. A stout femur and tibia are described. The morphology of the femur is unique for early tetrapods, with fibular and tibial condyles of similar length and lacking an adductor crest. The morphology of the femur, and its length relative to the tibia, suggests that the holotype of I. bradleyae preserves an immature individual. The tibia is a flat bone characteristic of stem tetrapods. Phalanges from the right and left pes are present. Because the phalanges are disarticulated, a phalangeal count cannot be determined. Despite the new anatomical information, the systematic position of I. bradleyae is still difficult to establish; however, it does not belong within the colosteids, temnospondyls, or embolomeres, to which it has previously been assigned.

Upper Ordovician strata exposed from the Baiyanhuashan section is the most representative Late Ordovician unit in the northwestern margin of the North China Craton (NCC). In total, 1,215 conodont specimens were obtained from 24 samples through the Wulanhudong and Baiyanhuashan formations at the Baiyanhuashan section. Thirty-six species belonging to 17 genera, including Tasmanognathus coronatus new species, are present. Based on this material, three conodont biozones—the Belodina confluens Biozone, the Yaoxianognathus neimengguensis Biozone, and the Yaoxianognathus yaoxianensis Biozone—have been documented, suggesting that the Baiyanhuashan conodont fauna has a stratigraphic range spanning the early to middle Katian. The Baiyanhuashan conodont fauna includes species both endemic to North China and widespread in tropical zones, allowing a reassessment of the previous correlations of the Katian conodont zonal successions proposed for North China with those established for shallow-water carbonate platforms at low latitudes.

UUID: http://zoobank.org/7cedbd4a-4f7a-4be6-912f-a27fd041b586

A new wood-boring ichnospecies is described from transgressive (lagoonal) deposits of the Lower Cretaceous Sparky Formation (Mannville Group) in west-central Saskatchewan, Canada. Apectoichnus lignummasticans new ichnospecies is a trace fossil that occurs in a thin coal bed and that was emplaced in an in situ xylic substratum (woodground). The ichnofossil is thin, elongate, unbranched, and straight to gently curved with a circular cross section and uniform diameter. Apectoichnus lignummasticans n. isp. is similar in many respects to modern borings in wood that are produced by marine isopods, e.g., Limnoria lignorum Rathke, 1799, for feeding and refugia. The recognition of Apectoichnus lignummasticans n. isp. in the rock record aligns with the modern observation that fossilized wood-boring assemblages should display higher ichnofossil diversities than commonly reported. Additionally, the stratigraphic occurrence of Apectoichnus lignummasticans n. isp. in association with other evidence of marine deposition reaffirms that certain wood boring morphologies (i.e., ichnotaxa) are useful as indicators of marine transgressions.

UUID: http://zoobank.org/880e722f-8944-42d7-bc38-423cc5a46413

The sparse record of Cretaceous crocodyliforms in Australia comprises only three species, all within the genus Isisfordia. Isisfordia duncani Salisbury et al., 2006 is from the Albian–Turonian Winton Formation of Queensland, and both Isisfordia molnari Hart et al., 2019 and Isisfordia selaslophensis Etheridge, 1917 have been described from opalized material from the Cenomanian Griman Creek Formation of New South Wales. Here, we describe new cranial and postcranial material, including the most complete crocodyliform skeleton from the Cretaceous of New South Wales, which is assigned to Isisfordia cf. I. selaslophensis. We also reappraise previously described crocodyliform material from the same locality. We find that much of this material displays features that are consistent with Isisfordia.

Metoposaurids are a widespread and ubiquitous constituent of Late Triassic non-marine paleoenvironments. In North America, this group is practically the only large-bodied temnospondyl clade, and is particularly well documented from the American southwest and south-central regions (Arizona, New Mexico, Texas). However, metoposaurids are poorly documented from eastern North America, with fragmentary, doubtfully diagnostic historical material such as “Dictyocephalus elegans” Leidy, 1856 and “Eupelor durus” Cope, 1866. The Zions View (early Norian?) locality in Pennsylvania preserves more-complete material, which previous workers noted as belonging to “Buettneria perfecta” Case, 1922 (=Anaschisma browni Branson, 1905). However, the material has never been described in a fashion that characterizes the anatomy or that justifies the taxonomic assignment, yet it would represent the most complete material in eastern North America and a substantial expansion of this taxon's geographic range. Here we redescribe the Zions View metoposaurid material in detail, differentiating it from Calamops paludosus Sinclair, 1917, the only other Late Triassic temnospondyl from the eastern seaboard, and demonstrating confident affinities with A. browni. Our study is the first to properly justify the taxonomic referral, underscoring the broader importance of proper documentation of voucher specimens, especially for potential geographic outliers. Anaschisma browni is thus the most widely dispersed metoposaurid. Its easternmost documentation underscores the importance of the undersampled and understudied metoposaurid record on the eastern seaboard for understanding the development of a metoposaurid zone of exclusivity in North America and demonstrates the need for further exploration to refine conceptualizations of Late Triassic tetrapod evolution.

A new genus and species of carneyellid edrioasteroid, Spiracarneyella florencei n. gen. n. sp., is described from the Upper Ordovician (Kaitian) Point Pleasant Formation of northern Kentucky and southern Ohio. Spiracarneyella n. gen. is characterized by having all five ambulacra curving clockwise around the theca, having small node-bearing interambulacral plates in the distal interambulacra, and having the periproct placement slightly offset to the right side of the CD interambulacrum. The oral area of carneyellids evolved by paedomorphosis of the oral plates covering the mouth. The straight ambulacra of Cryptogoleus and the spiraling ambulacra of Spiracarneyella n. gen. evolved by paedomorphosis and peramorphosis, respectively.

UUID: http://zoobank.org/79733c8f-0bc8-4e7e-8f77-8508f576755c

Polykampton recurvum n. isp. is the sixth ichnospecies of the ichnogenus Polykampton Ooster, 1869. It is a horizontal structure composed of a median cylindrical tunnel and narrow, usually back-curved lateral lobes located in alternating position. It occurs 2–3 cm below the top of single beds in the Maastrichtian–Paleocene deep-sea turbiditic marlstones of the Monte Antola Unit in the Northern Apennines. The lobes of P. recurvum n. isp. are actively filled with gray mudstone from above through the permanently open median tunnel. The trace fossil belongs to the category sequestrichnia, which is typical of oxygenated deep-sea environments characterized by seasonal or episodic supply of organic matter into a generally oligotrophic environment. P. recurvum n. isp. was produced by a “worm,” probably a polychaete, which adapted to seasonal or only episodic supply of organic matter to the deep-sea floor. The tracemaker stored the organic-rich mud in the lobes for nutrition during times of low organic matter availability on the seafloor.

UUID: http://zoobank.org/49555117-1c39-4658-8b49-3f3e2f47ba27

Herbivory is a common ecological function among extant lepidosaurs, but little is known about the origin of this feeding strategy within Lepidosauria. Here we describe a sphenodontian (Lepidosauria) from the Late Triassic of western North America, Trullidens purgatorii n. gen. n. sp., that reveals new aspects of the earliest radiation of herbivorous lepidosaurs. This taxon is represented by an isolated lower jaw with robust structure bearing transversely widened dentition and extensive wear facets, suggesting a masticatory apparatus specialized for herbivory. An unusual ‘incisor-like’ tooth is present at the anterior end of the jaw; a unique feature among lepidosaurs, this tooth is convergent with the incisors of extant rodents and lagomorphs. Phylogenetic analyses support the placement of this taxon within opisthodontian sphenodontians, a group sharing derived cranio-dental morphologies specialized for herbivory. The new taxon was recovered in a recently discovered and unnamed series of Upper Triassic strata in southeastern Colorado, USA, exposed in Canyons incised by the Purgatoire River and its tributaries. These strata comprise a dominantly red-bed sequence of conglomerates, sandstones, and siltstones deposited in a fluvio-lacustrine setting, preserving a Late Triassic biota of invertebrate and vertebrate ichnofossils, plant macrofossils, bony fish, temnospondyl amphibians, and reptiles. We use aetosaur osteoderms as biostratigraphic links to the nearby Chinle Formation of Arizona, USA, establishing a middle Norian age for these strata. The presence of an opisthodontian from western equatorial Pangaea in the Norian Stage reveals a near-global radiation of this clade across the Pangaean supercontinent during the Late Triassic.

UUID: http://zoobank.org/A737c03f-863a-488e-a860-5cc914548774.

Although the fossil record of the Late Cretaceous eastern North American landmass Appalachia is poor compared to that from the American West, it includes material from surprisingly aberrant terrestrial vertebrates that may represent relictual forms persisting in relative isolation until the end of the Mesozoic. One intriguing question is to what extent eastern and western North American faunas interspersed following the closure of the Western Interior Seaway during the Maastrichtian Stage of the Late Cretaceous ca. 70 Ma. Isolated remains from the Atlantic Coastal Plain in New Jersey have been preliminarily identified as the bones of crested lambeosaurine hadrosaurids, a derived clade known from the Cretaceous of Asia, western North America, and Europe, but have not been formally described. We describe the partial forelimb of a large hadrosaurid from the late Maastrichtian New Egypt Formation of New Jersey. The ulna preserves multiple deep scores identifiable as shark feeding marks, and both bones show ovoid and circular marks attributable to invertebrates. This forelimb is very similar to another partial antebrachium from the same area that shows evidence of septic arthritis. Both these specimens and a complete humerus from the same unit are closely comparable to the lower forelimbs of lambeosaurines among hadrosaurid dinosaurs. Although the absence of lambeosaurine synapomorphies observable on the New Egypt Formation forelimbs precludes their definite referral to Lambeosaurinae, they show that a morphotype of large hadrosauromorph with distinctly elongate forelimbs existed in the latest Maastrichtian of eastern North America and allow for a revision of the latest Cretaceous biogeography of crested herbivorous dinosaurs.

The family †Peipiaosteidae contains the genera †Peipiaosteus, †Stichopterus, †Spherosteus, †Yanosteus, and †Liaosteus, all from Late Jurassic to Early Cretaceous deposits of China, Russia, Kazakhstan, and Mongolia. Although the family has taxonomically expanded since it was first established for †P. pani Liu and Zhou, 1965, the amount of detailed comparative data for many of the taxa involved is lacking. In this paper, we describe the osteology of the monotypic genus †Yanosteus from the Yixian Formation (Early Cretaceous) of China largely on the basis of a newly prepared, well-preserved specimen. †Yanosteus is characterized by a series of infraorbital ossicles (a characteristic of the family), a broad, rounded palatopterygoid, a robust dentary, an extremely small opercle and a subopercle with distinctly long and rounded anterior process and a posteriorly scalloped margin, a broad and weakly forked caudal fin, an elongate dorsal fin with more than 160–178 fin rays (diagnostic for the genus), and a short but well-formed pectoral fin spine. We use the results of this study to discuss the characters of the †Peipiaosteioidei and the diversity of †peipiaosteioids.

The dispersal of Crocodylus from Africa to Europe during the Miocene is not well understood. A small collection of cranial fragments and postcranial elements from the latest Miocene (6.2 Ma) site of Venta del Moro (Valencia, Spain) have previously been referred to Crocodylus cf. C. checchiai Maccagno, 1947 without accompanying descriptions. Here we describe and figure for the first time the crocodylian remains from Venta del Moro, which represent at least two individuals. Our comparisons indicate that this material clearly does not belong to Diplocynodon or Tomistoma—the only two other crocodylians described so far for the European late Miocene. The material is only tentatively referred to cf. Crocodylus sp. because the apomorphies of this genus are not preserved and a referral to C. checchiai cannot be supported on a morphological basis. However, it is likely that this late Miocene species, originally described from Libya (As Sahabi) and later identified also in Kenya, could have dispersed across the Mediterranean Basin multiple times and colonized the southern areas of Mediterranean Europe, as evidenced by several Crocodylus or Crocodylus-like remains described during the past years.

The late Paleozoic temnospondyl Sclerocephalus formed an aquatic top predator in various central European lakes of the late Carboniferous and early Permian. Despite hundreds of specimens spanning a wide range of sizes, knowledge of the endocranium (braincase and palatoquadrate) remained very insufficient in Sclerocephalus and other stereospondylomorphs because even large skulls had unossified endocrania. A new specimen from a stratigraphically ancient deposit at St. Wendel in southwestern Germany is recognized as representing a new taxon, S. concordiae new species, and reveals a completely ossified endocranium. The sphenethmoid was completely ossified from the basisphenoid to the anterior ethmoid region, co-ossified with the parasphenoid, and the basipterygoid joint was fully established. The pterygoid bears a slender, S-shaped epipterygoid, which formed a robust pillar lateral to the braincase. The massive stapes was firmly sutured to the parasphenoid. In the temnospondyl endocranium, character evolution involved various changes in the epipterygoid region, which evolved distinct morphologies in each of the major clades.

UUID: http://zoobank.org/5e6d2078-eacf-4467-84cf-a12efcae7c0b

The new fossil micromammal assemblage of Çapak represents a mixture of both Anatolian and European faunal elements. The locality is very important for understanding faunal evolution in the less well-known time interval at the end of the early Miocene of western Anatolia. In Çapak, nine species of rodents and one species of ochotonid were encountered: the hamsters Democricetodon gracilis, Megacricetodon primitivus, Eumyarion aff. E. montanus, Cricetodon cf. C. aliveriensis, Cricetodon sp., and Karydomys cf. K. strati, the mole-rat Debruijnia sp., the squirrel Aliveria luteyni, the dormouse Myomimus tanjuae n. sp., and the pika Albertona balkanica. The assemblage is referable to Anatolian local zone E or MN unit 4. The relative abundance of the various genera is markedly different from that of the older early Miocene assemblages, suggesting that the environment in Anatolia became drier and had a more open landscape.

UUID: http://zoobank.org/75f3276c-dcd8-4090-b2f6-d8fc8d3c3f7c

Traversodontidae is a group of Triassic herbivorous/omnivorous cynodonts that represents the most diversified lineage within Cynognathia. In southern Brazil, a rich fossil record of late Middle/mid-Late Triassic cynodonts has been documented, with Exaeretodon riograndensis Abdala, Barberena, and Dornelles, 2002 and Siriusgnathus niemeyerorum Pavanatto et al., 2018 representing two abundant and well-documented traversodontids. The present study provides a comparative analysis of the morphology of the nasal cavity, nasal recesses, nasolacrimal duct, and maxillary canals of both species using computed tomography, highlighting the changes that occurred in parallel to the origin of mammaliaforms. Our results show that there were no ossified turbinals or a cribriform plate delimiting the posterior end of the nasal cavity, suggesting these structures were probably cartilaginous as in nonmammaliaform cynodonts. Both species show lateral ridges on the internal surface of the roof of the nasal cavity, but the median ridge for the attachment of a nasal septum is absent. Exaeretodon riograndensis and S. niemeyerorum show recesses on the dorsal region of the nasal cavity, which increase the volume of the nasal cavity, potentially enhancing the olfactory chamber and contributing to the sense of smell. On the lateral sides of the nasal cavity, the analyzed taxa show a well-developed maxillary recess. Although E. riograndensis and S. niemeyerorum have roughly similar nasal cavities, in the former taxon, the space between the left and right dorsal recesses of the nasal cavity is uniform along its entire extension, whereas this space narrows posteriorly in S. niemeyerorum. Finally, the nasolacrimal duct of S. niemeyerorum is more inclined anteroposteriorly than in E. riograndensis.

The highly fossiliferous St. Marys Formation is exposed along Calvert Cliffs, Maryland, and comprises one of the best available records of late Miocene life in the northeastern United States. Rachycentron stremphaencus new species, a cobia from the late Miocene (Tortonian) of the St. Marys Formation is described herein on the basis of a single three-dimensional neurocranium. This fossil represents the earliest known occurrence of neurocranial remains of the genus Rachycentron in the record. Rachycentron stremphaencus differs from Rachycentron canadum (Linnaeus, 1766) in many ways. The most obvious include a different ornamentation of the outer surface of the cranial bones; a notably pronounced lateral ridge resulting in a considerable gradient from the dorsal-medial exposure of the frontal to its lateralmost supraorbital margin; the size, shape, and position of the sphenotic that is located in the posterior half of the neurocranium and its lateralmost edge being adjacent to the anteriormost extent of the wedge-shaped trough in the dorsal surface of the skull formed by the lateral and medial ridges; the two contralateral medial ridges forming a proportionately much wider trough on either side of the supraoccipital; the epioccipitals not reaching the rear edge of the neurocranium; and the lack of a conspicuous posterolateral prong of the intercalar.

UUID: http://zoobank.org/c8523bb4-2de2-4318-9a37-027f463a0441

The Upper Triassic Tiki Formation of India has yielded several new cynodont taxa, which are described on the basis of multiple isolated teeth and a jaw fragment. A new species of dromatheriid, Rewaconodon indicus, is defined by a tri- and tetracuspid asymmetric crown, long anterior edge of the major cusp a, cingular cusps d and f, and marked constriction at the crown-root junction. Another new dromatheriid, Inditherium floris n. gen. n. sp., is characterized by a broad, flower-shaped pentacuspid crown, multiple cingular cusps, and a weak lingual cingulum is also described from the same horizon. In addition, a new mammaliamorph taxon, Tikiodon cromptoni n. gen. n. sp., is established on a tooth specimen, which has a shovel-shaped crown, three closely spaced main cusps, a pronounced lingual cingulum with multiple cingular cusps, and a root of incomplete root bifurcation. Such a tooth morphology occupies an intermediate position between the non-mammalian cynodonts and the early mammals, as is evident from the co-occurrence of various cynodont dental morphotypes in the Tiki Formation. Moreover, Late Triassic cynodonts occurred along narrow belts demarcated by paleolatitudes, though the Indian fauna shows both Laurasian and Gondwanan affinities.

UUID: http://zoobank.org/c2c575ad-ee23-4f33-8a30-661c548a5b17

Studies focused on deciduous dentition, ontogenetic series, and tooth eruption and replacement patterns in fossil mammals have lately increased due to the recognized taxonomic and phylogenetic weight of these aspects. A study of the deciduous and permanent dentition of Interatherium and Protypotherium (Interatheriinae) is presented, based mainly on unpublished materials. Deciduous cheek teeth are brachydont and placed covering the apex of the respective permanent tooth; in addition, some morphological and metrical differences are observed along the crown height. Five dental ontogenetic stages are distinguished among the juvenile specimens on the basis of the degree of wear, the replacement of the deciduous premolars, and the eruption of the molars. The crown height and the wear degree of different Interatheriinae taxa show: (1) eruption pattern of molars in an anterior–posterior direction (M/m1 to M/m3); (2) pattern of replacement of deciduous premolars and eruption of permanent premolars in a posterior–anterior direction (dP/dp4 to dP/dp2 and P/p4 to P/p2); and (3) eruption of M/m3 before the replacement of dP/dp4. Results allow evaluating the diagnostic dental characteristics used to describe some interatheriines, as well as reinterpreting some taxonomic assumptions: the holotype of Protypotherium diversidens Ameghino, 1891 is recognized as a juvenile of another species of the genus, and the species is not validated, considering it as Protypotherium sp.; the holotype of Eudiastatus lingulatus Ameghino, 1891 falls in the variability of Protypotherium, becoming P. lingulatus new combination, tentatively maintaining the species and implying the synonymy between Eudiastatus and Protypotherium; and the holotype of Eopachyrucos ranchoverdensis Reguero, Ubilla, and Perea, 2003 is reinterpreted as bearing deciduous premolars.

The Cincinnatian (Katian) of the Cincinnati Tri-State area is widely regarded as one of the most fossiliferous sections known (Meyer and Davis, 2009). Echinoderms from these strata include well-described asteroids, crinoids, cyclocystoids, edrioasteroids, glyptocystoids, mitrates, and ophiuroids. John Pope discovered a partially articulated echinoderm in float from the Fairview Formation that does not correspond to any known Cincinnatian echinoderm. Although mentioned in Ubaghs (1966, as a presumable personal communication from Pope, 1960), Haude and Langenstrassen (1976), Reich (2001), and Reich and Haude (2004), this specimen at the Cincinnati Museum Center (CMCPIP 51316) has neither been described nor illustrated; yet, these authors attributed it to Volchovia Hecker, 1938 in the Class Ophiocistioidea. Questions swirl around this fossil: what is its complete morphology; does it belong to Volchovia; whether or not it can be assigned to Volchovia, is it an ophiocistioid? The first step to understand this enigmatic echinoderm is to illustrate and describe the specimen, which is the objective of this note.

A small temnospondyl skull from the upper Carboniferous Allegheny Group of Five Points, Ohio, is referred to a new dissorophoid, Palodromeus bairdi n. gen n. sp. The complete skull with mandibles is preserved in counterparts. It is characterized by, (1) elongated slit-like choana; (2) postfrontal, postorbital, and supratemporal bearing a distinct ridge; and (3) reduced parietal only two-thirds the length of the frontal. Phylogenetic analysis confirms a dichotomy between amphibamiforms and olsoniforms and places Palodromeus bairdi n. gen. n. sp. at the base of the olsoniforms, revealing an early stage in the evolution of that clade.

UUID: http://zoobank.org/3871e443-7888-4439-a7c1-9463bc4d87ab