Titanopterans are spectacular, giant, predatory insects mainly known from the Triassic, but they are known from a few localities in Central Asia (including European Russia) and Australia. The Nampo Group is a nonmarine sequence, located on the southwestern Korean Peninsula, the age of which has remained controversial, due to lack of proper age-constraining fossils. Here, we report a new titanopteran Magnatitan jongheoni n. gen. n. sp. from the Amisan Formation, Nampo Group. The division of the radius anterior and radius posterior beyond the distal half of the fossil wing is a characteristic of the family Paratitanidae. This is the first discovery of a titanopteran fossil outside of Central Asia and Australia, suggesting a possible circum-Tethys oceanic distribution. Given the possibly widespread distribution of titanopterans, this group might have played a critical role as giant predatory insects in Triassic terrestrial ecology. Because titanopterans have never been found beyond the Triassic-Jurassic boundary, the occurrence of a titanopteran corroborates the Triassic age of the Nampo Group.

UUID: http://zoobank.org/10177029-0044-424f-9816-e3fb057cfe85

The beetle Ceutorhynchus is used as a proxy for Eocene core Brassicaceae. The age of Brassicaceae has been strongly debated because their fossil record is scanty. There are four species of Ceutorhynchus inaffectatus species-group known in the Priabonian: Ceutorhynchus zerovae new species in Rovno amber and three in Baltic amber. There are numerous extant members of the group, all of which feed on core Brassicaceae. Together with differentiation of Brassicaceae-feeding Pierinae (Lepidoptera) in the Priabonian, the strong presence of Ceutorhynchus indicates at least an early Priabonian age of core Brassicaceae. The oldest fossil Brassicaceae is not known in the late Eocene of Europe, but was recorded in Montana, dated in some studies as late Oligocene, but recently as Priabonian (34 Ma). Ceutorhynchus zerovae n. sp. is very close to C. electrinus from Baltic amber.

UUID: http://zoobank.org/7f10761f-463d-44c5-9eef-bb4697bfb116.

Exceptionally preserved fossil eggs and embryos provide critical information regarding paleoembryogenesis, reproductive strategies, and the early ontogeny of early arthropods, but the rarity of preservation of both eggs and egg-bearing organisms in situ limits their use in detailed evolutionary developmental (evo-devo) studies. Burgess Shale-type deposits preserve rare instances of egg-bearing arthropods as carbonaceous compressions; however, the eggs are usually poorly preserved with no compelling evidence of embryos. We describe the first record of a brooding specimen of Waptia cf. W. fieldensis from the Spence Shale, a Cambrian (Wuliuan Stage) Burgess Shale-type deposit in northeastern Utah and southeastern Idaho. This is the first record of an egg-bearing arthropod from the Spence Shale and it exhibits two distinct modes of preservation among eggs within the single clutch: carbonization and phosphatization. Unlike the egg-bearing Burgess Shale specimens, many eggs of this Utah specimen are also preserved three-dimensionally. In addition, synchrotron radiation X-ray tomographic microscopy reveals internal distributions of mineral phases, along with potential remnants of the egg membrane and attachment structures, but, as in the Burgess Shale, no explicit traces of developing embryos. The distinct modes of preservation highlight the existence of diagenetic microenvironments within some eggs, but not in others during fossilization.

A new species of drywood termite (Isoptera: Kalotermitidae) is described from a nearly complete alate specimen preserved in early Miocene Ethiopian amber. Glyptotermes abyssinicus new species is distinguished by its U-shaped head with 12-segmented antennae, the ocelli separated from the eye margin, the right mandible with an obtuse angle between the apical and first marginal teeth, the left mandible with an obtuse angle between the apical and first + second marginal teeth, and the wing venation. This is the first termite reported from Ethiopian amber, and the fourth Miocene species of the extant genus Glyptotermes Froggatt, 1897, together with species previously described from diatomites of China and amber from the Dominican Republic. As the oldest report of the genus known from Africa, G. abyssinicus n. sp. constitutes an interesting new record for the biogeographical history of the kalotermitid lineage.

UUID: http://zoobank.org/7670b045-fb31-4809-8116-4d14c4dd275b

Early “ptychoparioid” trilobites are likely to play a crucial role in determining the currently unresolved phylogenetic relationships among the numerous Cambrian libristomate clades. However, such phylogenetic analyses are hindered by the fact that many species and genera of early “ptychoparioids” are known from very limited material and are typically defined only on cranidial features. Herein, we report new “ptychoparioid” assemblages from the Saline Valley Tongue, Harkless Formation at Clayton Ridge, Nevada. These middle to upper Dyeran Stage (Cambrian Series 2, Stage 4) assemblages contain well-preserved early “ptychoparioids” that are represented by articulated exoskeletons as well as isolated cranidia, rostral plates, librigenae, thoracic segments, and pygidia, which together provide morphological details for the complete or nearly complete exoskeleton of three new genera and four new species: Anebocephalus silverpeakensis n. gen. n. sp.; Coenoides scholteni n. gen. n. sp.; Harklessaspis rasettii n. gen. n. sp.; and H. parvigranulosus n. gen. n. sp. Pending a formal phylogenetic analysis, it is deemed preferable to assign these taxa to new genera rather than to shoehorn them in to potentially ill-diagnosed existing genera. These new genera are compared to the Cambrian Series 2 taxa Crassifimbra and Eokochaspis, and the Miaolingian Series taxon Elrathina, which are similar in morphology and known from sclerites other than just cranidia.

Also documented in this study are the “ptychoparioids” Crassifimbra walcotti, Cr.? sp. A, Cr. sp. B, and “ptychoparioid” spp. A, B, C, and D from the Saline Valley Tongue and the overlying Mule Spring Limestone and lowermost Emigrant Formation.

UUID: http://zoobank.org/864d4005-2676-4f3f-b676-911efde7d263

The Devonian saw the emergence and subsequent diversification of the clam shrimp. To date, there are no credible records of any clam shrimp prior to the Devonian. Therefore, discoveries and taxonomic studies of early forms, especially the Early Devonian species, are essential to broaden our knowledge of the origin and early evolution of clam shrimp. An Early Devonian clam shrimp community that consists of four species, Cornia cheni n. sp., Pseudestheria cf. P. diensti, Palaeolimnadiopsis zhangi n. sp., and ?Palaeolimnadia sp. is described and discussed in detail in this study. These species are the oldest clam shrimp in China and among the oldest worldwide. This discovery indicates that clam shrimp communities with high diversity existed since the late Early Devonian and that their early forms may have had a profound effect on the origin and evolutionary trends of the following forms.

UUID: http://zoobank.org/ae9d0ab8-0b0e-4103-ad71-601d11a024fc.

The extinct family Hylicellidae, as the ancestral group of modern cicadomorphans, had a wide distribution and a very high species-level biodiversity from the Triassic to Early Cretaceous. We herein report 11 new hylicellid specimens from the Jurassic Daohugou beds of Inner Mongolia, NE China, and execute geometric morphometric analysis (GMA) to elucidate their systematic position. Our GMA and subsequent morphometric statistics indicate that 10 of our new specimens can be compared to the holotype of Cycloscytina gobiensis, and one is new to science. Cycloscytina incompleta new species is erected based on this specimen, with the following discriminatory tegminal traits: C3 almost as long as and slightly narrower than C2, and the forking position of stem M distinctly migrates towards wing apex and much apicad of the stem CuA bifurcating. Additionally, Cycloscytina plachutai is herein transferred to the procercopid genus Procercopina, resulting in P. plachutai new combination. To date, just a few body structures of Hylicellidae have been revealed, and the new whole-bodied hylicellids reported herein provide some novel insights on the evolution of basal Clypeata. This study also emphasizes the use of morphometric analysis in the systematics of wing-bearing insects such as hylicellids.

UUID: http://zoobank.org/84a67eba-9b7c-4e27-a436-764802c4cdfb.

Temnospondyl amphibians are common in non-marine Triassic assemblages, including in the Fremouw Formation (Lower to Middle Triassic) of Antarctica. Temnospondyls were among the first tetrapods to be collected from Antarctica, but their record from the lower Fremouw Formation has long been tenuous. One taxon, ‘Austrobrachyops jenseni,’ is represented by a type specimen comprising only a partial pterygoid, which is now thought to belong to a dicynodont. A second taxon, ‘Cryobatrachus kitchingi,’ is represented by a type specimen comprising a nearly complete skull, but the specimen is only exposed ventrally, and uncertainty over its ontogenetic maturity and some aspects of its anatomy has led it to be designated as a nomen dubium by previous workers. Here, we redescribe the holotype of ‘C. kitchingi,’ an undertaking that is augmented by tomographic analysis. Most of the original interpretations and reconstructions cannot be substantiated, and some are clearly erroneous. Although originally classified as a lydekkerinid, the purported lydekkerinid characteristics are shown to be unfounded or no longer diagnostic for the family. We instead identify numerous features shared with highly immature capitosaurs, a large-bodied clade documented in the upper Fremouw Formation of Antarctica and elsewhere in the Lower Triassic. Additionally, we describe a newly collected partial skull from the lower Fremouw Formation that represents a relatively mature, small-bodied individual, which we provisionally refer to Lydekkerinidae; this specimen represents the most confident identification of a lydekkerinid from Antarctica to date.

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.

Records of abnormal fossil arthropods present important insight into how extinct forms responded to traumatic damage and developmental complications. Trilobites, bearing biomineralized dorsal exoskeletons, have arguably the most well-documented record of abnormalities spanning the Cambrian through the end-Permian. As such, new records of malformed, often injured, trilobites are occasionally identified. To further expand the documentation of abnormal specimens, we describe malformed specimens of Lyriaspis sigillum Whitehouse, 1939, Zacanthoides sp. indet., Asaphiscus wheeleri Meek, 1873, Elrathia kingii (Meek, 1870), and Ogygiocarella debuchii (Brongniart, 1822) from lower Paleozoic deposits. In considering these forms, we propose that they illustrate examples of injuries, and that the majority of these injuries reflect failed predation. We also considered the origin of injuries impacting singular segments, suggesting that these could reflect predation, self-induced damage, or intraspecific interactions during soft-shelled stages. Continued examination of lower Paleozoic trilobite injuries will further the understanding of how trilobites functioned as prey and elucidate how disparate trilobite groups recovered from failed attacks.

Two new genera and six new species of trilobites are systematically documented herein: Sinagnostus mirabilis new genus new species, Yanpingia punctata n. gen. n. sp., Illaenus taoyuanensis n. sp., Panderia striolatus n. sp., Nileus yichongqiaoensis n. sp., and Paratiresias peculiaris n. sp. The materials were collected from the Darriwilian (late Middle Ordovician) strata in the Upper Yangtze Region, South China. Also provided is an emended diagnosis of the genus Paratiresias based on the new species Paratiresias peculiaris, which is the oldest known species of this genus with an extremely narrow (sag. and exsag.) preglabellar field. Those Chinese species previously referred to Nanillaenus are reassigned to Illaenus sensu lato. These trilobites add new data for the Darriwilian trilobite macroevolution and show highly endemic to South China and the faunal exchanges between South China and Tarim, Kazakhstan, Alborz, as well as Sibumasu and North China.

UUID: http://zoobank.org/ec3be9be-b003-4367-910d-7a0ac4edc982

Despite the abundant literature on limnic Cretaceous ostracode faunas, the database on mid-Late Cretaceous taxa is still scarce. The Songliao Basin in northeastern China preserves a diverse assemblage of ostracode fossils from the Santonian–Campanian Nenjiang and Sifangtai formations. This rather unique material is of major importance to comprehend Early to mid-Late Cretaceous (Cenomanian–Campanian) limnic ostracode faunas and therefore the evolution of Late Cretaceous basins in China. A study of this fauna was conducted to detail the taxonomy of ostracode species from members 1 and 2 of the Nenjiang and Sifangtai formations in the Songliao Basin. Well-preserved specimens of 14 ostracode species were recovered from samples of the ZKY2-1 well in southwestern Songliao Basin: Scabriculocypris liaukhenensis Liu in Netchaeva et al., 1959, Ilyocyprimorpha netchaevae Su in Netchaeva et al., 1959, Cypridea acclinia Netchaeva in Netchaeva et al., 1959, Cypridea cavernosa Galeeva, 1955, Cypridea gracile Netchaeva in Netchaeva et al., 1959, Cypridea lepida Ye in DOFEAD, 1976, Cypridea squalida Sou in Netchaeva et al., 1959, Fabaeformiscandona? Disjuncta (Hao in Hao et al., 1974), Lycopterocypris profunda Lübimova, 1956, Mongolocypris magna (Hou, 1958), Mongolocypris tera (Su in Netchaeva et al., 1959), Talicypridea obliquecostae (Szczechura and Błaszyk, 1970), Talicypridea reticulata (Szczechura, 1978) and Renicypris renalata (Su in Hao et al., 1974). Four of the species identified received emended descriptions and diagnoses. Other important results include the first discussion on sexual dimorphism in Cypridea acclinia and Fabaeformiscandona? disjuncta, the first study on the ontogeny of Ilyocyprimorpha netchaevae, and the reassignment of Cypridea liaukhenensis to the genus Scabriculocypris. Several species recorded herein are also found in other continental far-eastern Asian basins, while genera range from worldwide to far-eastern Asian distribution; these results support strong affinities among faunas of Mongolia, China, and Japan.

Predation is of great importance in understanding the diversification of spatangoid echinoids as well as their preservation in the fossil record. Herein, the spatangoid-dominated Miocene carbonate succession of Punta Foghe (Montiferru area, central-western Sardinia, Italy) is studied with the aim of reconstructing paleoenvironmental conditions and investigating abiotic and biotic factors, including predation, that influence the paleoecology and preservation potential of spatangoid echinoids.

The echinoid fauna is dominated by the spatangoid Agassizia and, to a lesser extent, the clypeasteroid Clypeaster. The fauna also includes both regular (undetermined diadematoids and camarodonts) and irregular (Plagiobrissus, Echinocyamus, and Koehleraster) forms. Echinoids and associated fossil content, along with taphonomic and sedimentological signatures, indicate an inner sublittoral environment with moderate energy conditions affected by high-energy events. Agassizia remains, which consist mainly of complete and well-preserved tests, commonly bear subcircular drill holes that are interpreted as the result of cassid gastropod predation and investigated with respect to size selectivity and stereotypy of attack sites. Potential biases related to drilling predation and biostratinomy on the preservation potential of spatangoid tests are discussed.

Agassizia lacks any morphological adaptation to minimize high predation risk, including defensive spines and the ability to burrow deeply into the sediment. An antipredator behavior possibly relied on a gregarious life-history strategy, reducing the frequency of attacks on specific individuals by cassid predators.

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.

Sloths were among the most diverse groups of land vertebrates that inhabited the Greater Antilles until their extinction in the middle-late Holocene following the arrival of humans to the islands. Although the fossil record of the group is well known from Quaternary deposits in Cuba, Hispaniola, and Puerto Rico, remains from older units are scarce, limiting our understanding of their evolution and biogeographic history. Here we report the oldest known fossil ground sloth from Hispaniola, represented by an unassociated partial tibia and scapula that are recognized as a single taxon from the late Miocene-early Pliocene of the Dominican Republic. The combination of characters observed on the tibia suggests a close relationship with Megalocnus, otherwise only known from the Pleistocene–Holocene of Cuba. These fossils fill a temporal gap between those previously known from the early Miocene of Cuba and those from Pleistocene–Holocene deposits in the region and provide additional support for a continuous presence of the group in the Greater Antilles since the Oligocene.

Two cupressocrinitids (Crinoidea, Eucladida) from the famous crinoid collection of Dr. Ludwig J.T. Schultze deposited in the Museum of Comparative Zoology (MCZ) are revised in the present study: Cupressocrinites hybridus n. comb. and Halocrinites minor n. comb. The rare C. hybridus has a stratigraphically and regionally restricted occurrence at the Eifelian–Givetian boundary of the Hillesheim Syncline (Eifel, Rhenish Massif, Germany), but the more common H. minor occurs supraregionally from the Eifelian to the Frasnian (Middle to Upper Devonian). Both cupressocrinitids are redescribed on the basis of the type material stored in the MCZ. In addition, the rare Halocrinites heinorum n. sp. from the Eifelian–Givetian boundary beds and lower Givetian of the Eifel (Rhenish Massif, Germany) is described, and Encrinus townsendi (König, 1825) is questionably reassigned to Robustocrinites: R.(?) townsendi n. comb. Pre- and postmortem ossicular modifications of the studied cupressocrinitid skeletons are also discussed.

UUID: http://zoobank.org/4598e1fa-8ca5-4160-8fee-6e8c6d444115

The Ladinian–Carnian transition in the Tethys domain was accompanied by an important environmental change representing a milestone in the climate evolution of the Triassic. However, estimations on paleodiversity composition and paleoenvironmental conditions across this interval are scarce in marine settings due to the lack of fossil-bearing successions. In this work, a refined paleontological and sedimentological study has allowed us to better characterize a well-preserved marine ?Ladinian–Carnian carbonate succession in the South Central Pyrenees (Odèn site, Catalonia, NE Spain). Vertebrate faunas include numerous actinopterygian specimens, forming an assemblage composed of at least four taxa: Peltopleurus cf. P. nuptialis Lombardo, 1999, Saurichthys sp., Colobodus giganteus (Beltan, 1972), and an indeterminate halecomorph. Specimens belonging to the genus Peltopleurus are dominant; the long-snouted Saurichthys, the halecomorph, and the large-bodied Colobodus giganteus are less abundant. Tetrapod remains are scarcely present and are assigned to sauropterygians. Invertebrate faunas include bivalves (Pseudocorbula gregaria [Münster in Goldfuss, 1838]) and brachiopods (Lingula sp.). The fossil assemblage was recovered from organic-rich laminated silty mudstone layers. Sedimentological and textural analyses suggest that fossil biotas were deposited below the fair-weather wave base in shallow subtidal coastal settings. These environments were sporadically sourced by silt/clay. The age of the Odèn site, on the basis of the recovered fauna, is assigned to the ?late Ladinian–middle Carnian (Middle–Late Triassic), which is in agreement with previously published ages based on palynomorph data.

The refined integration of paleontological, sedimentological, and biostratigraphic data from the Odèn site and other vertebrate-bearing localities in the Tethys domain can help better constrain the paleoenvironmental conditions and paleogeographical configuration impacting ecosystem diversity during the late Ladinian–Carnian interval.

A new specimen of the arachnid order Trigonotarbida is described from the Middle Pennsylvanian (lower Desmoinesian) Shelburn Formation of Indiana, which has previously yielded the remains of a phalangiotarbid. Two new trigonotarbid arachnid specimens are also described from the Middle Pennsylvanian (Desmoinesian) Senora Formation of Oklahoma. These are the first trigonotarbid specimens reported from Indiana and Oklahoma. The Indiana trigonotarbid belongs to the Eophrynidae, as indicated by distinct features such as the large tubercles on the dorsal surface of the opisthosoma and two pairs of terminal opisthosomal spines. This specimen is the first arachnid fossil to be imaged using a Multistripe Laser Triangulation scanner. The heavy dorsal tuberculation, lobed and subtriangular carapace, rounded clypeus, lack of terminal opisthosomal spines, and rounded opisthosoma on Oklahoma specimen FMNH PE 56932 indicate it belongs to the genus Aphantomartus, in Aphantomartidae. The other Oklahoma specimen, FMNH PE 56955, possesses opisthosomal tergites that are divided into five plates longitudinally as well as a subquadrate carapace, which identify it as a member of Anthracomartidae; its rounded opisthosomal margin shows it to belong to the genus Anthracomartus.

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.

UUID: http://zoobank.org/48dcbe31-e25d-4657-8441-732d5ea2b49d

Based on abundant topotype specimens from the Oil Creek Formation at the type locality in southern Oklahoma, Histiodella labiosa is described and revised. The morphological variation of the Pa, Pb, and M elements of this previously little-known species is also systematically documented and illustrated. The revised species definition, with formal designation of a lectotype and a new diagnosis supported by morphological comparison with its ancestral and descendent species, has enhanced its value for correlation of the lower Darriwilian. The inferred phylogeny of Histiodella that resulted from this study is based on both cladistic and morphological analyses. It supports the monophyletic grouping of Histiodella species and their evolutionary relationships. This has wide biostratigraphic implications because many Histiodella species are valuable for detailed global correlation of upper Dapingian to middle Darriwilian strata.