The Cambrian (Miaolingian; Wuliuan) Spence Shale Lagerstätte of northern Utah and southern Idaho is one of the most diverse Burgess Shale-type deposits of Laurentia. It yields a diverse fauna consisting of abundant biomineralized and locally abundant soft-bodied fossils, along a range of environments from shallow-water carbonates to deep-shelf dark shales. Panarthropods are the dominant component throughout the deposit, both in time and space, but whereas the trilobites and agnostoids are abundant, most of the soft-bodied taxa are only known from very few specimens. Additionally, the knowledge of soft-bodied panarthropods is currently largely limited to locations in the Wellsville Mountains of northeastern Utah. This contribution describes 21 new soft-bodied panarthropods from six locations, including the first occurrences of soft-bodied panarthropods in the High-Creek, Smithfield Creek, Spence Gulch, and Two-Mile Canyon localities. Additionally, we report the presence of bradoriids— i.e., Branchiocaris pretiosa Resser, 1929, Perspicaris? dilatus Robison and Richards, 1981, Naraoia? sp. indet., Thelxiope cf. T. palaeothalassia Simonetta and Delle Cave, 1975, and Tuzoia guntheri Robison and Richards, 1981—for the first time from the Spence Shale Lagerstätte; the first reported occurrence outside of the Burgess Shale for Thelxiope cf. T. palaeothalassia; and the first Wuliuan occurrence of Tuzoia guntheri. We also report on a new hurdiid carapace element and additional specimens of Buccaspinea cooperi? Pates et al., 2021, Dioxycaris argenta Walcott, 1886, Hurdia sp. indet., and Tuzoia retifera Walcott, 1912. This new material improves our understanding of the panarthropod fauna of the Spence Shale Lagerstätte and substantially increases our understanding of the distribution of the described taxa in time and space.

Although knowledge of their fossil record continues to improve, multituberculates nonetheless remain one of the more poorly understood mammalian clades, which can be attributed to a record comprised of isolated teeth and fragmentary jaws. Fortunately, the p4 of multituberculates is the most common form of remains for this group and is a principal source of diagnostic characters in systematic studies, the p4 of cimolodontan multituberculates is both common and a source of diagnostic characters in systematic studies. The results of a recent morphometric study on the neoplagiaulacid Mesodma suggest that p4 size may be more useful than shape in diagnosing the various species referred to this genus. We tested this hypothesis by applying two different morphometric methods (2D geometric morphometrics and linear measurements) to two samples: (1) one including the p4s of four known species of Mesodma (M. ambigua, M. thompsoni, M. formosa, and M. pygmaea), and (2) a sample of unidentified p4s of Mesodma from the Bug Creek Anthills locality of northeastern Montana. Our results indicate that while form explains most of the morphological variation in p4s of the various species of Mesodma, linear-measurement data support differences in p4 morphology that are not recovered by form data alone. Depending on the methods used, we found evidence for the presence of one or more species of Mesodma in the Bug Creek Anthills fauna. Although shape and size both contribute to morphological variation in the p4 of Mesodma, our results suggest that the diagnostic power of each varies with the type of methodology employed.

The suborder Phacopina, characterized by schizochroal eyes, is among the most common groups of trilobites in Devonian strata. The marine sediments of the Famennian in western Junggar, Xinjiang, contain abundant low-disparity phacopids, which have previously been designated to Omegops accipitrinus mobilis, Phacops circumspectans tuberculosus, and Omegops cornelius on the basis of small numbers of poorly preserved specimens. In this study, these phacopids were identified as two new species of Omegops, O. honggulelengensis n. sp. and O. xiangi n. sp., on the basis of nearly 200 well-preserved specimens. The intraspecific variations of eye lenses of these specimens were quantitatively analyzed. On the basis of differences in the total number, number of dorsoventral files, and arrangement of the eye lenses, the absence of lenses in the middle part of the visual surface, and asymmetry of the number and/or arrangement of lenses in the two eyes, it was concluded that the reasons for intraspecific variation in eye lenses of Late Devonian Omegops from western Junggar were different from previously described factors but were likely genetic or embryological malfunctions or abnormalities caused by pathological conditions. Diversity of lenses in the schizochroal eyes shows that the number and arrangement of eye lenses was not stable in Phacopina. Therefore, many specimens are needed for quantitative study to determine the true characteristics of the number or arrangement of eye lenses when these features are used in the systematic taxonomy of Phacopina.

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The first fossil Typhlocybinae inclusions from Eocene Rovno amber are described and illustrated. They include two new monotypic genera of the extinct tribe Protodikraneurini, Retrorsotettix n. gen. with type species R. vlaskini n. sp. and Protoparallaxis n. gen. with type species P. clavatus n. sp. Also described and illustrated are two new monotypic genera of the extant tribe Dikraneurini, Eodikraneura n. gen. with type species E. obscura n. sp. and Rovnodikra n. gen. with type species Rovnodikra longipes n. sp. Retrorsotettix is the oldest leafhopper known to exhibit a false eyespot and false leg markings on the forewing, representing an early acquisition of a defensive strategy against visual predators. Appearance of a small insect with false eyespots in the Eocene fossil record may reflect increased pressure by visual predators, especially crown ornithuromorph insectivorous birds. Such birds and small insect prey with false eyespots remain unknown in the Cretaceous. Eodikraneura exhibits a unique condition of the hind wing venation in which radius posterior and media anterior veins are confluent for a short distance but then diverge before separately reaching the submarginal vein. This presumably represents a morphological transition between the Protodikraneurini, in which hind-wing radius posterior and media anterior veins are completely separate and connected by a radial-medial crossvein, and Dikraneurini, in which radius posterior and media anterior veins are completely confluent distally. A key to genera of Protodikraneurini is provided.

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This work describes a rhynchonellid and spire-bearing brachiopod fauna from the Ixtaltepec Formation of Oaxaca, Mexico. Leiorhynchoidea perrilliatae, Allorhynchus scientiana, and Anthracospirifer oaxacaensis are new species. The specific determination, along with information of previously described taxa, allowed the establishment of precise relative ages of the different fossiliferous intervals (API-1 to API-8) of the formation. The occurrence of Serpukhovian taxa in API-1 to API-3 allowed assignment of the strata to the Upper Mississippian. The presence of Bashkirian species allowed the assignment of the rocks of API-5 and API-6 to the Lower Pennsylvanian. Likewise, Middle Pennsylvanian brachiopods in API-7 and API-8 enabled correlation of the strata with the Moscovian stage. This study shows that the Ixtaltepec Formation is represented by a succession of well-delimited Serpukhovian, Bashkirian, and Moscovian rocks. Regarding paleogeography, the brachiopod fauna displays clear taxonomic variations that concur with global geological changes that occurred between the Serpukhovian to Moscovian. In the Serpukhovian intervals, we recorded numerous cosmopolitan taxa of tropical waters, coinciding with the migration pathway of the Rheic Ocean. For the Bashkirian, we observed a North American provincialism; however, because of the presence of Australian and South American species, it is proposed that the Austropanthalassic-Rheic corridor had a close connection with Oaxaca. The main provincialism was observed in the Moscovian association because most of those taxa have been reported from different localities in the United States. This study supports that the main resemblance between Oaxacan and North American faunas continued until the Pennsylvanian and not the Mississippian, as was previously proposed.

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The decapod crustaceans of Central European Russia have been the subject of studies since the nineteenth century, and the only species, Eryma quadriverrucatum Trautschold, 1866 (Erymidae), has been found in the Callovian to the Oxfordian of that region. The present paper discusses the new exceptional findings of Solenoceridae, Glypheidae, and Mecochiridae from the upper Callovian sites of the Ryazan Region. The previously reported presence of Archeosolenocera sp. is confirmed in detail; Glypheopsis aff. G. etalloni (Oppel, 1861) and Eumorphia sp. are noticed for the first time for this area. Modern identification of erymids is reported: Eryma aff. E. ventrosum (von Meyer, 1835), E. ornatum (Quenstedt, 1858), and Stenodactylina insignis (Oppel, 1862) are noted in addition to E. quadriverrucatum. These findings address the gaps in the fragmented knowledge about the decapod fauna of Central European Russia and exhibit a differentiation of the local Russian decapod assemblage from those of Western Europe. The decapod community of the Ryazan region is considered to be associated with soft substrates of upper sublittoral zone.

An early form of eurypterids (Chelicerata), Archopterus anjiensis n. gen. n. sp., is described from the uppermost Ordovician Wenchang Formation of Anji County, Zhejiang Province, South China. It is the earliest record of eurypterids in China and likely the oldest representative of the Adelophthalmidae. The species, represented by a single specimen, is diagnosed by a ventrally preserved prosoma with a parabolic carapace, Hughmilleria-type prosomal appendages, a short vase-shaped metastoma, and a three-segmented type A (female) genital appendage. The specimen is preserved, together with diverse sponges, graptolites and occasional nautiloids, in a 10 m thick shale of marine deep-water setting. This finding extends the stratigraphic range of adelophthalmids from the previously recorded early Silurian into the Late Ordovician (some 10 million years older) and supports an earlier cryptic phase of eurypterid evolution in Gondwana.

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We report new material of the rare early Eocene rodent Lophiparamys debequensis Wood, 1962 from the Willwood Formation of the southern Bighorn Basin, north-central Wyoming. The new material constitutes the first record of L. debequensis from the Bighorn Basin and documents aspects of the anatomy of Lophiparamys that were previously unknown, including a portion of the maxilla and a portion of the tarsus. The maxillary fragment demonstrates that Lophiparamys has a small P3 and a relatively large infraorbital canal. The tarsus of Lophiparamys is similar to that of other early rodents but differs in a few features that suggest an arboreal locomotor repertoire, including an asymmetric astragalar trochlea, long astragalar neck, transverse astragalar sustentacular facet, short calcaneal tuber, elongate calcaneal ectal facet, and circular calcaneal cuboid facet. The presence of arboreally adapted features in the tarsus of Lophiparamys is consistent with a hypothesized relationship between small-bodied Eocene microparamyine rodents and extant Gliridae. Phylogenetic analysis fails to consistently support this relationship or monophyly of Microparamyinae, but both remain plausible. Comparison of L. debequensis with other species of the genus emphasizes the distinctiveness of L. debequensis and suggests the presence of multiple lineages of Lophiparamys.

Zygocycloides? foerstei n. sp. is described from the Llandovery (Aeronian) Brassfield Formation of southwestern Ohio. This is among the oldest reported Silurian cyclocystoids from North America and is the only North American Llandovery cyclocystoid that is preserved with a complete rim of marginal ossicles. Zygocycloides Smith and Paul, 1982 is most similar to Nicholsodiscus Glass et al., 2003 (Katian) and Perforocycloides Ewin et al., 2019 (Llandovery, Telychian), both from Anticosti Island, Québec. Cyclocystoids (Ordovician to Mississippian) survived Late Ordovician extinctions, and this discovery documents that this echinoderm clade was part of shallow-water, marine paleocommunities during the initial post-extinction transgression onto the Laurentian platform.

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This work provides the detailed investigation (taxonomy, biostratigraphy, and paleoecology) of a nonmarine ostracod fauna from the Shinekhudag Formation in Gobi–Altai area, southwest Mongolia. The samples from two sections (Tsagaan Tsuvarga and Oshih Hollow East) yielded various ostracods assigned to nine species belonging to six genera: Cypridea verrucata Neustrueva, 1974, C. ihsienensis Hou, 1958, C. unicostata Galeeva, 1955, C. tumefacta Neustrueva, 1974, Yumenia cf. Y. oriformis Hou, 1958, Scabriculocypris subscalara Zhang and Chen in Ye et al., 2003, ?Trapezoidella sp., Candona sp., and Vlakomia ulanense Neustrueva, 1977. The biostratigraphic application of our taxonomic results suggests that the age of the Shinekhudag Formation is Barremian–Aptian. In addition, the ostracod fauna shows strong affinities to faunas from northeast China and potentially northwest China, providing evidence of faunal exchanges between these regions and southwest Mongolia. The new discovery of Vlakomia ulanense indicates that mid-Cretaceous (Albian) species in northeast China probably originated in Mongolia at the time of deposition of the Shinekhudag Formation. Two ostracod assemblages in the studied sections reflect that different paleoenvironment settings have existed during deposition; the Tsagaan Tsuvarga Section represents a shallow-littoral zone of athalassic (inland) saline lake under arid climate, and the dominance of cypridoideans of the Oshih Hollow East Section is indicative of ephemeral water bodies.

A diverse Wuchiapingian brachiopod fauna, which contains 57 species in 28 genera, is described from the Shuizhutang Formation at the Liannan section, Guangdong province, southeastern China. Four new species Tyloplecta liannanensis n. sp., Linoproductus huananensis n. sp., Araxathyris minor n. sp., and Permophricodothyris flata n. sp. are proposed. From well-preserved Liannan specimens, characteristics of the shell microstructures in Permianella are revised, and different morphologies of muscle scars in Permophricodothyris are distinctly shown. Until now, only several Wuchiapingian brachiopod faunas have been found in South China. Compared with these faunas, the Liannan fauna shows much higher α diversity and is more like faunas from southeastern China than those from the Yangtze area in faunal composition. The Liannan fauna is dominated with Neochonetes, Transennatia, Orthothetina, Permophricodothyris, and Cathaysia, which are normally larger and more strongly ornamented than their Changhsingian counterparts. The Wuchiapingian brachiopods in South China are represented mainly by the Douling fauna and Shuizhutang fauna. The Douling fauna has relatively low diversity and presents the survival stage after the Guadalupian–Lopingian boundary crisis. The Shuizhutang fauna has a much higher diversity and more key Changhsingian taxa and shows a rapid radiation stage. Faunal compositions of the two faunas indicate that the initial recovery of brachiopods occurred mainly at the genus level followed by a more rapid radiation at both genus and species levels.

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Among the several evolutionary lineages of the baleen whales (Mysticeti), the eomysticetids are an ancient successful family that retain possibly nonfunctional teeth and functional baleen, a transitional stage between toothed and baleen-assisted filter-feeding mysticetes. The patchy fossil record leaves gaps in eomysticetid paleobiology interpretations, but their diversity and widespread geographical distribution can be a relevant proxy to understanding the evolution of crown Mysticeti. Here we describe a new baleen whale, Echericetus novellus n. gen. n. sp., from the Oligocene of Mexico (slightly older than 27.95 million years ago). This new taxon has morphological features that show its affinity to Eomysticetidae, such as the intertemporal region longer than wide, the elongate and oval temporal fossa, and a well-developed and lobate coronoid process of the mandible. Similarly, our cladistic analyses confirm the inclusion of Echericetus in the Eomysticetidae. Echericetus reinforces our notion of the eomysticetid diversity and disparity. Geographically, the existence of Echericetus from Mexico also indicates that eomysticetid inhabited subtropical regions in the Northern Hemisphere. Last, our discovery of a new eomysticetid from the Oligocene of Mexico provides new insights into the distribution patterns and habitat use of Eomysticetidae, essential to further explain the demise of this transitional lineage between toothed and baleen-bearing whales.

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Crinoids were major constituents of late Carboniferous (Pennsylvanian) marine ecosystems, but their rapid disarticulation rates after death result in few well-preserved specimens, limiting the study of their growth. This is amplified for cladids, who had among the highest disarticulation rates of all Paleozoic crinoids due to the relatively loose suturing of the calyx plates. However, Erisocrinus typus Meek and Worthen, 1865 has been found in unusually large numbers, most preserved as cups but some as nearly complete crowns, in the Barnsdall Formation in Oklahoma. The Barnsdall Formation, a Koncentrat Lagerstätte, is composed predominantly of fine- to medium-grained sandstone, overlain by mudstone and shale; severe compaction of the fossils in the mudstone and shale layer in this formation allowed for exceptional preservation of the plates. Herein, we summarize a growth study based on 10 crowns of E. typus, showcasing a well-defined growth series of this species from the Barnsdall Formation, including fossils from juvenile stages of development, which are rarely preserved. We used high-resolution photographs imported into ImageJ and recorded measurements of the cup and arms for all nondistorted or disarticulated plates. Results show that the plates of the cup grew anisometrically with both positive and negative allometry. The primibrachial plates of E. typus grew with positive allometry. The brachial plates started as uniserial (i.e., cuneiform) as juveniles but shifted to be biserial. Erisocrinus typus broadly shares similar growth trajectories with other cladids. These growth patterns provide insight into feeding strategies and can aid in understanding crinoid evolutionary paleoecological trends.

A diverse echinoderm fauna lived in reef and non-reef Silurian facies of the upper Midwestern USA. However, these faunas are dominantly preserved in dolostones with moldic preservation, and fossils from dolostone facies have not been documented to the extent of Silurian crinoids in nondolostone strata. Herein, an echinoderm fauna is described from the dolostones of the Cedarville Member of the Laurel Limestone (Wenlock, Homerian) from the Pepcon Cement Quarry in west-central Ohio. The described fauna contains blastoids, hemicosmitoids, and crinoids, including Troosticrinus subcylindricus (Hall and Whitfield, 1875); Caryocrinites sp. indet.; an unidentifiable diplobathrid camerate; Periechocrinus tennesseensis (Hall and Whitfield, 1875); Periechocrinus egani? (Miller, 1881); Stiptocrinus farringtoni (Slocom, 1908); Calliocrinus primibrachialis Busch, 1943; Calliocrinus poepplemani new species; Calliocrinus hadros new species; and Lecanocrinus sp. indet. Generic concepts for the Eucalyptocrinitidae are clarified; and, surprisingly, Eucalyptocrinites Goldfuss, 1831 is absent from this fauna. Additionally, lectotypes and paralectotypes are designated for Periechocrinus tennesseensis and Calliocrinus primibrachialis.

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Pleurocystitid echinoderms are known to have consistent plating in the theca or body. However, many specimens with aberrant plating are known among glyptocystitoids (Glyptocystella, Hadrocystis, Tyrridiocystis, and so on) as well as the new Ordovician species Pleurocystites? scylla, described here from the Benbolt Formation near Thorn Hill, Tennessee, USA. The presence of an extra plate in the holotype of P? scylla is most likely teratological. While teratology is often misconstrued as parasitism and pathology, it is easily recognized among glyptocystitoid rhombiferans because of the conservative nature of plate arrangements found in these taxa. Teratologic features in stemmed echinoderm thecae appear only to alter suture positioning and not significantly change the thecal shape or diminish the diagnosability of a specimen taxonomically.

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The early geologic history of family Pinnidae, a diverse and abundant component of mud-dominated shallow marine environments of the late Paleozoic, is obscured by ill-defined genus and species concepts. Mistakes in descriptions and reconstructions have provided a basis for suggestions that older taxa in the family differ from younger taxa and may not belong in family Pinnidae. This report provides improved documentation that removes the basis for the concept of divergence. Late Paleozoic genera and species have definite pinnid bivalve characters, including a dorsomarginal fold that holds the ligament, a thick outer columnar prismatic shell layer, and an equivalved, triangular shell, all of which provide evidence for confident assignment of the genus to family Pinnidae. The suggested synonymy of Aviculopinna with Pteronites is invalid, and genus Aviculopinna is limited to occurrence in Permian strata. The two genera belong in different families. A neotype is designated for genus Aviculopinna type species Aviculopinna pinnaeformis Geinitz from Gera, Germany, and a lectotype for Aviculopinna neukirchensis Langenhan from Nowy Kościół, Poland. Inferences on the life habits of Aviculopinna based on its occurrence in Poland are presented. An evaluation of the subterminal beak versus terminal beak concept in pinnids is presented and conclusion presented that there are few data available to support the concept or to test it. In its present form, the concept has no utility in the study of pinnid bivalves.

Goryeocrinus pentagrammos n. gen. n. sp. from the Jigunsan Formation of South Korea is described and is the first diplobathrid recorded from Middle Ordovician (middle Darriwilian) strata of East Gondwana. Phylogenetic analyses suggest that G. pentagrammos n. gen. n. sp. is a member of the paraphyletic Rhodocrinitidae of the Diplobathrida and most closely related to Paradiabolocrinus from the Late Ordovician (Sandbian) of Laurentia. Goryeocrinus pentagrammos n. gen. n. sp. is characterized by having a pentameral, flat bowl-shaped calyx, a conspicuous pentagrammatic ridge formed by bifurcated median ray ridge and pentagonal basal ridge, at least two interradials in the first row of regular interrays, an anitaxial ridge originating from the CD interray but close to the C ray radial, and lacking intrabrachials and interradials between secundibrachials. The occurrence of G. pentagrammos n. gen. n. sp. from South Korea (East Gondwana) drastically expands the Ordovician paleogeographic range of camerates, which otherwise have been recorded from Laurentia, West Gondwana, Avalonia, and Baltica.

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The typical Mississippian shallow-water deposits of the Jiusi and Shangsi formations are well exposed in the Yashui section in southern Guizhou, South China. The strata are composed mainly of platform limestones intercalated with shales and sandstones. Conodonts obtained from the limestones are dominated by two assemblages of cavusgnathids: the Cavusgnathus aff. Cav. unicornis and Clydagnathus windsorensis assemblage from the upper Jiusi Formation and the Cavusgnathus unicornis assemblage from the lowest Shangsi Formation. Vogelgnathus campbelli (Rexroad, 1957) is reported for the first time in Eastern Paleotethys and is characterized by small lateral denticles. The evolutionary first occurrence of Cavusgnathus unicornis Youngquist and Miller, 1949 is recognized as a marker for the regional Shangsian stage. This biostratigraphic datum is interpreted to be correlative with other shallow-water conodont zones marked by Cavusgnathus unicornis s.l. and similar species from North America and Europe.

Evidence from the earliest-known crinoids (Tremadocian, Early Ordovician), called protocrinoids, is used to hypothesize initial steps by which elements of the calyx evolved. Protocrinoid calyces are composed of extraxial primary and surrounding secondary plates (both of which have epispires along their sutures) that are unlike those of more crownward fossil and extant crinoids in which equivalent calycinal plating is strongly organized. These reductions inspired several schemes by which to name the plates in these calyces. However, the primary-secondary systems seen in protocrinoids first appeared among Cambrian stem radial echinoderms, with primaries representing centers around which secondaries were sequentially added during ontogeny. Therefore, the protocrinoid calyx represents an intermediate condition between earliest echinoderms and crownward crinoids. Position and ontogeny indicate certain primaries remained as loss of secondaries occurred, resulting in abutting of primaries into the conjoined alternating circlets characteristic of crinoids. This transformative event included suppression of secondary plating and modification or, more commonly, elimination of respiratory structures. These data indicate subradial calyx plate terminology does not correspond with most common usage, but rather, supports an alternative redefinition of these traditional expressions. Extension and adoral growth of fixed rays during calyx ontogeny preceded conjoined primaries in earliest crinoids. Restriction with modification or elimination of calyx respiratory structures also accompanied this modification. Phylogenetic analyses strongly support crinoid origination from early pentaradiate echinoderms, separate from blastozoans. Accordingly, all Tremadocian crinoids express a distinctive aggregate of plesiomorphic and apomorphic commonalities; all branch early within the crinoid clade, separate from traditional subclass-level clades. Nevertheless, each taxon within this assemblage expresses at least one diagnostic apomorphy of camerate, cladid, or disparid clades.