New fossil insect specimens from two new localities in Germany, namely Sperbersbach and Cabarz (Goldlauter Formation; early Permian), belonging to the Grylloblattida, are described. Abundant material is assigned to Pictoborella clara n. gen. n. sp., regarded as closely related to Pictoborella germanica (Prokop et al., 2012) n. comb., from the Saar-Nahe Basin (Germany; early Permian). Liomopterum fuscatum n. sp., represented by fewer specimens, is delimited based on previously published and new data on various Liomopterum spp. Two other Liomopteridae, Uralioma thuringiensis n. sp. and Liomopterites sperbersbachensis n. sp., each known from a single forewing, are also described. Finally, Cabarzopterum magnificus n. gen. n. sp., with unclear familial affinities, is described based on three forewings. The assemblages of grylloblattidan insects at Sperbersbach and Cabarz are generally similar, with differences probably related to different depositional environments and local paleoclimatic conditions.

UUID: http://zoobank.org/25ae125c-6a17-415f-8cf0-4c0507e4f89d

Bryozoans are common and diverse in fossil and modern coral reefs. However, studies of bryozoans in ancient reefs are generally limited, and even less is known about fossil bryozoan faunas associated with coral reefs in the Caribbean region. This is the second contribution describing the bryozoan assemblage from the early Miocene (Aquitanian) reefs of the Siamaná Formation in the La Guajira Peninsula, southern Caribbean. Here, we describe and illustrate 17 species of ascophoran-grade cheilostomes, including one new genus and three new species, Ditaxiporina colombiana n. sp., Poricella paulae n. sp., and Cycloavicularia parva n. gen. n. sp. Of the remaining fourteen taxa left in open nomenclature, one is considered confer and three affinis to species previously described, one is identified at family level, and nine at genus level. The Siamaná bryozoan fauna differs in species and colony-form composition from those associated with other paleoenvironments from Oligocene and Miocene localities of North America, the Caribbean, and Brazil.

UUID: http://zoobank.org/043eadcf-0e39-4c1b-b207-f7628d6b5c84

Borings and bite marks on fossil turtle carapaces and plastra from the Miocene Moghra Formation, northern Egypt, are herein described. All fossil turtle material from Moghra exhibits ichnofossils. The positions of invertebrate borings on external surfaces of tortoise and turtle shell material at Moghra are consistent with the activities of ectoparasites or mesoparasites.

A single invertebrate ichnotaxon, Karethraichnus lakkos Zonneveld et al., 2016, occurs on Moghra tortoise fossils. This trace fossil was likely emplaced by ixodid arthropods (ticks). Bite marks assigned to Nihilichnus occur on a carapace peripheral and are interpreted to reflect postmortem scavenging. An abundant and moderately diverse assemblage of invertebrate borings characterizes Moghra aquatic turtle shells. Karethraichnus lakkos and Thatchtelithichnus holmani Zonneveld et al., 2016 traces on aquatic turtles are interpreted to reflect leech and/or trematode parasitism. Gunnellichnus moghraensis (new ichnogenus new ichnospecies) and G. akolouthiste (n. isp.) likely reflect bacterial and/or fungal infections on aquatic turtle shells that rarely dried out.

UUID: http://zoobank.org/f0a3977d-deff-4954-b196-536adc405854

Records of diagenetically mineralized, filamentous, cavity-dwelling microorganisms extend back to strata from the early Paleoproterozoic (2400 Ma). In North Greenland (Laurentia), they are first known from the Ediacaran (Neoproterozoic; ca. 600 Ma) Portfjeld Formation of southern Peary Land, in association with a biota similar to that of the Doushantuo Formation of China. The Portfjeld Formation cavity dwellers are compared with more widespread occurrences in Cambrian (Series 2, Stage 4, Miaolingian Series) strata from the same region in which assemblages in postmortal shelter structures within articulated acrotretoid brachiopods and other invertebrates are common. All specimens were recovered by digestion of carbonate samples in weak acids. The described fossils are preserved as mineral encrusted threads but this diagenetic phosphatization unfortunately obscures their biological identity.

The fusuline genera Thailandina Toriyama and Kanmera, 1968 and Neothailandina Toriyama and Kanmera, 1968 were established by Toriyama and Kanmera (1968) based on material from the Khao Phlong Phrab section of the Permian Rat Buri Limestone in central Thailand that is currently assigned to the Khao Khad Formation of the Saraburi Group (Ueno and Charoentitirat, 2011). These fusuline genera are peculiar in having parachomata and replaced tests by secondary mineralization. Moreover, Neothailandina was described to have a test with transverse septula, considered to be characteristic for Neoschwagerinidae. Based on these remarkable test features, Toriyama and Kanmera (1968) newly introduced the subfamily Thailandininae to accommodate these two new genera and assigned it to the Neoschwagerinidae, despite the lack of septula in Thailandina. Later, Kobayashi et al. (2010) argued that Thailandina and Neothailandina are just a mixed grouping of several known genera of schwagerinids, verbeekinids, and neoschwagerinids that are too altered by recrystallization to be recognizable, and rejected the taxonomic validity of these two genera as well as Thailandininae.

The Khao Phlong Phrab section represents one of the standard late Cisuralian−Guadalupian (late early−middle Permian) fusuline successions in the eastern Paleotethys (Zhang and Wang, 2018) and contains not only Thailandina and Neothailandina but also abundant schwagerinid, verbeekinid, and neoschwagerinid fusulines (Toriyama, 1975; Fig. 1). I investigated the original specimens described by Toriyama and Kanmera (1968) and Toriyama (1975) from the section that are housed in the Department of Earth and Planetary Sciences of Kyushu University, Japan. I found that most of the grounds for Kobayashi et al.'s (2010) arguments to regard the thailandinin genera as taxonomically invalid are not supported by observations on these specimens as explained in the account that follows. In this taxonomic note, I propose that Thailandina and Neothailandina, and their family Thailandinidae, should be retained as valid taxonomic groups.

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 moderately diverse assemblage of brachiopods from the Latham Shale Lagerstätte (Cambrian Series 2, upper Stage 4) and the upper Cadiz Formation (Miaolingian, Wuliuan), California is described in detail for the first time. The fauna includes both linguliform and rhynchonelliform brachiopods—Hadrotreta primaea (Walcott, 1902), Paterina prospectensis (Walcott, 1884), Dictyonina pannula (White, 1874), and Mickwitzia occidens Walcott, 1908; and Nisusia fulleri Mount, 1981 and Wimanella highlandensis (Walcott, 1886), respectively—together with olenellid trilobites and hyolithids. The fauna differs from other Cambrian Konservat-Lagerstätten (notably Cambrian Series 2 Chengjiang and Guanshan Lagerstätten, eastern Yunnan) in that the brachiopod shell valves in many cases are still preserved with their original mineralization. Moreover, the excellently preserved shale-hosted valves even include cases with exquisite epithelial cell molds, otherwise only seen in acid-etched material from carbonate rocks. The pitted ornamentation in D. pannula closely resembles that described from Ordovician linguliforms. The unusual preservation of N. fulleri provides important clues for ancestral composition of the brachiopod shell. The two articulated rhynchonelliform species probably represent the oldest records of this group from the west Laurentia. The fauna could also represent the earliest onset of the transition from the Cambrian Evolutionary Fauna (CEF) to the Paleozoic Evolutionary Fauna (PEF).

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.

A diverse, well-preserved radiolarian assemblage is reported from the Sandbian age Climacograptus bicornis Graptolite Biozone. This new assemblage, recovered from the Pingliang Formation in the Guanzhuang section, China, includes six new species along with 13 other previously described taxa. Geminusphaera new genus incorporates G. grandis n. sp. and G. kongtongensis n. sp. and is proposed for inaniguttids constructed from two distinct porous spheres bearing seven or more primary spines. Protopylentonema new genus is introduced to incorporate pylomate entactinarians with five-rayed initial spicules. It includes P. ordosensis n. sp. as well as P. aperta, P. rimata, and P. insueta that were formerly assigned to Kalimnasphaera. Micro-computed tomography investigation of skeletal microstructure supports establishment of the new genera together with other new spumellarian species: Haplotaeniatum implexa n. sp., Inanigutta quadrispinosa n. sp., and Kalimnasphaera pingliangensis n. sp. It also indicates that family-level reassignment of Etymalbaillella from the Proventocitidae to the Ceratoikiscidae is appropriate.

Global distribution of Late Ordovician radiolarian occurrences highlights a strong preference for areas with equatorial to tropical sea surface temperatures. This must have led to ecological stress among radiolarian communities in adapting to global cooling in the Hirnantian.

UUID: http://zoobank.org/3d3f55b8-0e70-4f9f-9738-265750d8ec3a.

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.

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

A revision of the rhinocerotid material from the Negev (Israel), dating back to the early Miocene (MN3 in the European Mammal Biochronology), highlights the presence of Brachypotherium and a taxon close to Gaindatherium in the Levantine corridor. A juvenile mandible, investigated using CT scanning, displays morphologically distinct characters consistent with Brachypotherium cf. B. snowi rather than with other Eurasian representatives of this genus. Some postcranial remains from the Negev, such as a humerus, display features that distinguish it among Miocene taxa. We attribute these postcrania to cf. Gaindatherium sp., a taxon never recorded outside the Siwaliks until now. This taxon dispersed into the Levantine region during the late early Miocene, following a pattern similar to other South Asian taxa. Brachypotherium cf. B. snowi probably occurred in the Levantine region and then in North Africa during the early Miocene because its remains are known from slightly younger localities such as Moghara (Egypt) and Jebel Zelten (Libya). The occurrence cf. Gaindatherium sp. represents a previously unrecorded range expansion out of Southeast Asia. These new records demonstrate the paleogeographic importance of the Levantine region showcasing the complex role of the Levantine corridor in intercontinental dispersals between Asia and Europe as well as Eurasia and Africa.

In the first half of the nineteenth century, a marked shift occurred in our understanding and treatment of the chelicerate fossil record, with the differentiation and recognition of entirely extinct genera for the first time. At the heart of this taxonomic revolution were the Eurypterida (sea scorpions) and Xiphosura (horseshoe crabs), although both groups were in fact considered crustaceans until Lankester's (1881) seminal comparative anatomical study of the extant xiphosuran Limulus Müller, 1785 and modern scorpions. The oldest available eurypterid genus is Eurypterus deKay, 1825; the oldest available fossil arachnid genus name is that of the scorpion Cyclophthalmus Corda, 1835. However, there has been considerable historical confusion over the oldest available fossil xiphosuran genus name, which has been recognized alternately as Belinurus König (with a publication date of either 1820 or 1851) or the synonymous Bellinurus Pictet, 1846. Most recent treatments (e.g., Selden and Siveter, 1987; Anderson and Selden, 1997; Anderson et al., 1997; Lamsdell, 2016, 2021; Bicknell and Pates, 2020) have favored Bellinurus Pictet, 1846 as the available name; however, Haug and Haug (2020) recently argued that Belinurus König, 1820 is valid and has priority, a position then followed by Lamsdell (2020), prompting a reinvestigation of the taxonomic history of the genus. Upon review, it is clear that neither of the previously recognized authorities for Belinurus are accurate and that the two candidate type species for each genus are, in fact, synonyms. Given the convoluted and at times almost illogical history of the competing names, along with the most recent controversy as to which has priority, we present a complete history of the treatment of the genus to resolve the issue.

Olenellid trilobites from the lower Cambrian of the Iberian Peninsula are very scarce and poorly studied, making them difficult to compare with defined species and to include in biostratigraphic and paleobiogeographic analyses. Based on newly collected specimens, we revise the species ‘Callavia? lotzei’ Richter and Richter, 1941 from the ‘Cumbres beds’ of Cumbres de San Bartolomé and the ‘Herrerías shale’ of Cañaveral de León, Sierra del Bujo, and Hinojales (Huelva, Spain), and ‘Paradoxides choffati’ Delgado, 1904 from the Vila Boim Formation of Elvas (Portugal). The new material indicates that Callavia? lotzei is a junior synonym of ‘P. choffati.’ The Iberian species are here assigned to Callavia Matthew, 1897, for which morphological characters are reassessed, offering a valuable opportunity to discuss characters previously misinterpreted for this genus. Traditionally assigned to the Olenelloidea Walcott, 1890, Callavia lacks some of the diagnostic characters of this superfamily and is here assigned to Judomioidea Repina, 1979. A new diagnosis for this genus is provided, and Sdzuyomia Lieberman, 2001 is considered to represent a junior synonym of Callavia. The genus Callavia is distributed across the western margin of Gondwana, from the western Mediterranean region (Iberia and Morocco) throughout all the Avalonia sectors (UK, eastern Newfoundland, and Massachusetts). Its presence in Iberia supports the faunal links between the West Gondwana domain and Avalonia during Cambrian Series 2. The Iberian records of Callavia choffati are assigned to the middle part of the regional Marianian Stage (uppermost Cambrian Stage 3 to the lowermost Cambrian Stage 4) and correlates with the Callavia Biozone of Avalonia (lower Branchian Series).

Protandrous sex change (sex change from male to female) is one of the diverse sexual expressions exhibited by bivalves, but its expression in the shell is quite rare. Previous studies on living and fossil astartids suggest a relationship between protandrous sex change and the formation of crenulations on the ventral shell margin at later ontogenetic stages. Here we report the formation of such crenulations in the Early Jurassic astartid Nicaniella rakoveci (Kuhn, 1935) from the Amaltheenton Formation at Buttenheim, Franconia. This is the earliest known record of protandrous hermaphroditism in fossil bivalves, predating previous reports by at least 13 Myr. A principal component analysis of linear size measurements and Fourier shape analysis of the shell outlines revealed that the outline of Nicaniella rakoveci specimens varies from subquadrate to subcircular, but this variation is independent of the presence or absence of crenulations and therefore not associated with sex. Crenulated specimens exhibit a lower height/inflation ratio than non-crenulated ones, suggesting that the protandrous females have more inflated valves than the males. The formation of crenulations was probably related to allocation of resources for reproduction. The most likely function of the crenulations was to increase the internal shell volume in the female stage to accommodate more eggs rather than being an adaptation against predation as often assumed for other bivalves. The formation of crenulations is part of the protandrous life history and probably is controlled by a genetic mechanism that is also responsible for sex change.

Dragastanella transylvanica n. gen. n. sp. is described. Its calcified skeleton contains numerous voids, partly related to the molds of soft parts of the alga, but also related to lack of calcification. Interpretation of these voids, especially their attribution to original structures (e.g., primary lateral versus reproductive organ), has important implications for the taxonomic position of the alga, even at the family level. Examination of key sections that include the boundary between sterile and fertile parts of the alga excludes the occurrence of external reproductive organs. Unusual, paired pores in the outer part of the mineralized skeleton reflect an asymmetry within the whorl, excluding the presence of secondary laterals. The alga is characterized by a cylindrical to club-shaped thallus bearing only phloiophorous primary laterals arranged in whorls and flaring outwards, forming a cortex. Mineralized lenticular reproductive organs containing cysts set in the equatorial plane (Russoella-type gametophores) occur inside primary laterals (cladosporous arrangement of the reproductive organs). These characters support establishment of the new genus Dragastanella. Dragastanella transylvanica n. gen. n. sp. resembles species previously referred to Zittelina (Zittelina hispanica and Zittelina massei) and Triploporella (Triploporella matesina and Triploporella carpatica). Except for Triploporella carpatica, whose mineralized skeleton does not permit confident attribution to either Triploporella or Dragastanella n. gen., the other species must be ascribed to Dragastanella n. gen. Therefore, the following new combinations are proposed: Dragastanella hispanica n. comb., Dragastanella massei n. comb., and Dragastanella matesina n. comb. Despite widely overlapping biometrical measurements, these species can be differentiated by the size and location of their reproductive organs, the pattern of calcification around the primary laterals, and relationships among structural parameters such as the size of laterals, number of laterals per whorl, and distance between whorls.

The Early Ordovician (late Tremadocian; Stairsian) trilobite Gonioteloides Kobayashi has long been known from a small number of pygidia assigned to a single formally named species, and its affinities have not been assessed. Silicified material from western Utah and southeastern Idaho includes six distinct species assigned to the genus, one of which is the type species. Two others (G. moffitti and G. pankowskii) are new and formally named. An additional three species that are clearly new but known from sparse material are described in open nomenclature. Gonioteloides has a stratigraphic distribution through five consecutive trilobite zones in the mid-Stairsian Stage (upper Tremadocian). Although exoskeletal morphology of three species is almost completely known, the phylogenetic affinity of the taxon remains difficult to determine. It is tentatively assigned to Dimeropygidae Hupé.

UUID: http://zoobank.org/23257d6c-262b-4ef5-ae4e-cc431777e67e

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

An assemblage of 50 species of small shelly fossils is described from Cambrian Series 2 (Stage 4) strata in North Greenland, the present day northernmost part of the paleocontinent of Laurentia. The fossils are derived from the basal member of the Aftenstjernesø Formation at Navarana Fjord, northern Lauge Koch Land, a condensed unit that accumulated in a sediment-starved outer ramp setting in the transarctic Franklinian Basin, on the Innuitian margin of Laurentia. Most other small shelly fossil assemblages of similar age and composition from North America are described from the Iapetan margin of Laurentia, from North-East Greenland south to Pennsylvania. Trilobites are uncommon, but include Serrodiscus. The Australian bradoriid Spinospitella is represented by a complete shield. Obolella crassa is the only common brachiopod. Hyoliths, including Cassitella, Conotheca, Neogloborilus, and Triplicatella, are abundant and diverse, but most are represented just by opercula. Sclerites interpreted as stem-group aculiferans (sachitids) are conspicuous, including Qaleruaqia, the oldest described paleoloricate, Ocruranus?, Inughuitoconus n. gen., and Hippopharangites. Helcionelloid mollusks are diverse, but not common; they are associated with numerous specimens of the bivalve Pojetaia runnegari. The fauna compares best with that of the upper Bastion Formation of North-East Greenland, the Forteau Formation of western Newfoundland, and the Browns Pond Formation of New York, but several taxa have a world-wide distribution. Many specimens are encrusted with crystals of authigenic albite. New species: Anabarella? navaranae, Stenotheca? higginsi, Figurina? polaris, Hippopharangites groenlandicus, Inughuitoconus borealis, and Ocruranus? kangerluk.

UUID: http://zoobank.org/160a17b1-3166-4fcf-9849-a3cabd1e04a3