The lower-middle Hetang Formation (Cambrian Stage 2–3) deposited in slope-basinal facies in South China is well known for its preservation of the earliest articulated sponge fossils, providing an important taphonomic window into the Cambrian Explosion. However, the Hetang Formation also hosts a number of problematic animal fossils that have not been systematically described. This omission results in an incomplete picture of the Hetang biota and limits its contribution to the understanding of the early evolution of animals. Here we describe a new animal taxon, Cambrowania ovata Tang and Xiao, new genus new species, from the middle Hetang Formation in the Lantian area of southern Anhui Province, South China. Specimens are preserved as carbonaceous compressions, although some are secondarily mineralized. A comprehensive analysis using reflected light microscopy, scanning electron microscopy, energy-dispersive X-ray spectroscopy, and micro-CT reveals that the new species is characterized by a spheroidal to fusoidal truss-like structure consisting of rafter-like crossbars, some of which are secondarily baritized and may have been internally hollow. Some specimens have aperture-like structures that are broadly similar to oscula of sponges, whereas others show evidence of a medial split reminiscent of gaping carapaces. While the phylogenetic affinity of Cambrowania ovata Tang and Xiao, n. gen. n. sp. remains problematic, we propose that it may represent carapaces of bivalved arthropods or more likely sponges in early life stages. Along with other problematic metazoan fossils such as hyolithids and sphenothallids, Cambrowania ovata Tang and Xiao, n. gen. n. sp. adds to the diversity of the sponge-dominated Hetang biota in an early Cambrian deepwater slope-basinal environment.

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The morphology and growth habits of Evactinopora species of the Evactinoporidae (new family) are documented. This distinctive family of free-living bryozoans has a radial colony form at all growth stages. During a brief attachment phase on a hard substrate, the colony morphology grew as an expanding cone with vertical folds. Following detachment of the nascent colony from this hard substrate, it settled on soft sediment and the free-living expanding colony acquired a star-like form by producing slender outrigger rays. Continued growth produced a radial array of vertical vanes containing feeding autozooecia. The colony maintained a vertical orientation on soft sediment by means of outrigger rays and secretion of solid skeleton on the colony base that provided ballast. The radial growth pattern, outrigger rays, and vertical vanes developed as adaptive characters suitable for free-living life on soft sediment. North American species of Evactinopora are redefined and described taxonomically on the basis of zoarial and zooecial characters and a new species, Evactinopora mangeri, erected. The new family Evactinoporidae is established on the basis of the novel characters of early colony detachment from a hard surface, radial growth pattern through life, generation of outrigger rays, and growth of vertical vanes from the top of rays.

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The atrypide brachiopod Schachriomonia is documented here for the first time from the upper Katian Hadabulaktag Formation on the northeastern edge of Tarim Basin in Northwest China. These shells are conspecific with the earlier reported Rhynchotrema pentagonia Fang in Liu and Fang, 1990 from the same region. This species is most similar to Schachriomonia parva Popov et al., 1999 described from the Upper Ordovician of the Chu-Ili and Chingiz terranes and less closely resembles species from North and South China, indicating a close paleogeographic connection between these plates and terranes and Tarim during the Late Ordovician. Its dominance at three horizons in the study area may indicate that this species was an opportunistic colonizer of the seafloor following ecological disruption, or alternatively that the larvae were substrate-selective. Variation in shell shape from a nearly circular shell outline with a dorsal sulcus and ventral fold in the smallest shells to a wider outline with a more reversed dorsal fold and ventral sulcus in the largest is an example of ontogenic development. This may have been an adaptation to accommodate the growing lophophore or development of gonads in the mantle cavity over the life of the brachiopod. A stronger tooth-socket connection in larger shells strengthened the hinge area to better withstand the stresses caused by the mass of a larger shell.

The Chilean species traditionally assigned to the genera Chlamys Röding, 1798 or Zygochlamys Ihering, 1907 are now placed in two new endemic South American taxa: Dietotenhosen n. gen. (middle Miocene–early middle Pliocene), to include the southeastern Pacific Ocean species D. hupeanus (Philippi, 1887) n. comb. and D. remondi (Philippi, 1887) n. comb., and Ckaraosippur n. gen. (earliest middle Miocene–Pliocene), for C. calderensis (Möricke, 1896) n. comb. (Chile) and C. camachoi n. sp. (Argentina). Both genera are the youngest survivors of the tribe Chlamydini in southern South America. None of them is related to the circumpolar genus Psychrochlamys Jonkers, 2003, and the previous proposal of the dispersal through the Antarctic Circumpolar Current for the species included herein in Dietotenhosen is rejected.

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Major transitions in trilobite ontogeny have historically been defined based on the number and distribution of trunk segments, and articulation between the trunk and cephalon. This study documents additional morphological change across the meraspid-holaspid transition on the Ordovician phacopid trilobite Calyptaulax strasburgensis. An extensive dataset of silicified cranidia and pygidia collected from the mid-Ordovician Edinburg Formation of Virginia was subjected to a series of multivariate analyses, with a primary focus on the intersections and termini of furrows. Multivariate regression of partial warp scores demonstrates statistically significant change in allometric growth patterns over the course of development. These changes are concentrated in earlier instars, but are coincident in cranidia and pygidia. This sharp decrease in the rate of allometry, present in both tagmata, is expressed as significant breakpoints derived from a segmented regression, with the largest portion of allometric change found in the pre-breakpoint individuals. The term holeidos is proposed to describe the completion of form during trilobite development, independent of the completion of the thorax. The most dramatic change in shape during this period of ontogeny includes lateral glabellar expansion through deflection of the axial and palpebral furrows, possibly reflecting a change in the feeding habit during later development. Other morphological changes include the development of a more angular appearance to the anterior portion of the glabella, and anterior migration of the pygidial anterior margin. The appearance of these growth patterns in Calyptaulax extends the temporal range of these changes, some of which have only been documented in Devonian phacopids.

Although the order Odonata has a rich fossil record, many questions about its reproductive biology remain unanswered. There are two strategies of egg laying among odonates, exophytic and endophytic, the latter being one of the most revealing vestiges of plant–insect association in the fossil record. We assessed whether geometric morphometrics based on elliptical series of Fourier allow expression of variability of shape in traces of Odonata eggs within a leaf of Eucalyptus chubutensis (Berry) González (in part), González, 2009 (Myrtaceae) from Laguna del Hunco (Chubut, Argentina) (early Eocene) and whether this variability is consistent with the ichnotaxonomy of this material. We found that the largest variation corresponds to the compression of the shape while the remaining two components reflect variations in the apex position and its curvature, which changed according to the relative position of the traces in the leaf. There was no evidence that the hardness of the leaf would affect the shape of the egg trace. We postulate that these traces could have been produced by one single female: Variations in the pattern observable in the fossil of an originally three-dimensional structure are consistent with differences in the position of the eggs inserted by a single female who has flexed her abdomen to insert the eggs as she approaches the apex of the leaf (behavior observed also in extant dragonflies). For the first time, endophytic egg traces are analyzed with geometrical morphometrics, and this allows us to make inferences on the oviposition behavior of a female that lived around 52 million years ago.

End-Ordovician extinctions had a profound effect on shallow-water benthic communities, including the Crinoidea. Further, recovery after the extinctions resulted in a macroevolutionary turnover in crinoid faunas. Anticosti Island is the most complete Ordovician-Silurian boundary section recording shallow-water habitats. Both new taxa and changes in Anticosti Island stratigraphic nomenclature are addressed herein. New taxa include Becsciecrinus groulxi n. sp., Bucucrinus isotaloi n. sp., Jovacrinus clarki n. sp., Plicodendrocrinus petryki n. sp., Plicodendrocrinus martini n. sp., Thalamocrinus daoustae n. sp., and Lateranicrinus saintlaurenti n. gen. n. sp. The status of Xenocrinus rubus as a boundary-crossing taxon is confirmed, range extensions of several taxa are documented, and the distribution of crinoids with the revised stratigraphic nomenclature is documented.

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A new Lower Devonian fauna from the Iberian Chains (NE Spain) is described. Specimens have been collected from the shaley intervals of the Mariposas Formation dated as early Emsian. These include the camerates Acanthocrinus carsli n. sp., Platyhexacrinus santacruzensis n. sp., Culicocrinus breimeri n. sp., Camerata indeterminate, and an indeterminate eucladid. Compared with other faunas from Spain, this represents a low diversity crinoid assemblage that was probably concentrated in shallow, turbid environments. A summary of crinoids previously described from the Spanish Devonian is reported, which indicates that crinoid faunas become progressively more cosmopolitan during the Devonian.

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The Upper Ordovician Sandbian to Katian strata from the East Qilianshan (northeastern Tibet Plateau) bear a graptolite fauna of moderately high diversity. Graptolites from the Amplexograptus maxwelli beds to the Appendispinograptus longispinus Biozone (Sa2–Ka4 intervals) proposed herein include 27 species of 13 genera. This important graptolite fauna is first described herein although it was initially reported in 1963. Most of them occur in the A. longispinus Biozone corresponding to the Dicellograptus complexus to Paraorthograptus pacificus biozones of the Wufeng Formation in the Yangtze region. Alulagraptus new genus is established based on the materials from the East Qiqiaogou section. The endemic species, e.g., Alulagraptus ensiformis (Mu and Zhang in Mu et al., 1963), Dicellograptus sinicus Mu and Zhang in Mu et al., 1963, and Climacograptus? papilio Mu and Zhang in Mu et al., 1963, could indicate that East Qilianshan block was separated from South China.

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The base of the Emsian, which is defined by the first appearance of the conodont Polygnathus kitabicus, has never been successfully demonstrated in the South China Block (including Guangxi and eastern Yunnan). As a result, we studied conodonts from the lowermost part of the Shizhou Member of the Yukiang Formation at the Dacun-1 section in the Liujing area, Guangxi. This new investigation has revealed a conodont fauna only consisting of Polygnathus pireneae, P. sokolovi, P. kitabicus, P. sp. and Pandorinellina exigua philipi, which can be assigned to the uppermost part of the pireneae Zone and the lowermost part of the kitabicus Zone in ascending order. The Pragian/Emsian boundary at the Dacun-1 section is located in the highest thick-bedded limestone bed that can be observed in the lowermost part of the Shizhou Member. Therefore, this is the first time that the lower boundary of the Emsian defined by the lowest occurrence of P. kitabicus is reported in the South China Block. However, the scarcity of suitable limestone samples for conodont analysis in the middle and upper parts of the Shizhou Member precludes definitive identification of the upper boundary of the kitabicus Zone in the Liujing area.

The giant carnivorous phorusrhacid bird Phorusrhacos longissimus (Aves, Cariamiformes) was first described in 1887 by Florentino Ameghino on the basis of a jaw fragment. The majority of a skull of the species still encased in crumbling rock was preserved only long enough for illustrations to be made by Carlos Ameghino in the field and for a brief description to be written. Skull remains of this species have remained scarce, and few postcranial remains have been figured. Here, we reassess the cranial anatomy of this outstanding ‘terror bird’ species taking into account data from a newly discovered skull. An additional specimen of a well-preserved dorsal vertebra referable to Phorusrhacinae is also described from a separate locality within the Miocene Santa Cruz Formation (late early Miocene) from Santa Cruz Province in Argentina. The skull includes most of the rostrum, skull roof, and mandible and is compared with material from other members of the Phorusrhacinae. The new data from the skull and vertebra provide morphological features of this clade that benefit future taxonomic and phylogenetic analyses of this iconic group of birds.

Beremendiini is an extinct group of soricine shrews that were widely distributed during the Pliocene and Pleistocene. Their occurrence in China has been investigated, but their presence in northern Asian regions has remained poorly studied. This paper analyzes 56 fossil remains of Beremendiini collected from 16 early Pliocene to early Pleistocene localities in Russia (Siberia), Kazakhstan, and Mongolia and shows the presence of two beremendiin species: Beremendia fissidens (Petényi, 1864) and Beremendia minor Rzebik-Kowalska, 1976. Northern Asian Beremendia considerably vary in size and qualitative characteristics, although most of the different states have been identified in European or Chinese specimens. Through the application of geometric morphometric techniques, mandibular shape analyses reveal similarities between the members of the beremendiin genera Peisorex Kowalski and Li, 1963, Beremendia Kormos, 1934, and Lunanosorex Jin and Kawamura, 1996. Shape analyses and comparisons of mandibular characteristics reveal ‘trophic’ analogies between Beremendia and Blarina spp. and a new model of ‘Mandible Swinging and Sliding’ (MSS-model) accounting for the similarities in mandibular morphology with implications for the understanding of the diet of Beremendia.

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Xenarthra is an endemic South American lineage of mammals, probably the sister clade of the other placental mammals. The oldest records of Xenarthra are from the latest Paleocene, although its current diversity is much lower than that recorded in some intervals of the Cenozoic Era. A new Neogene Xenarthra (Pilosa and Cingulata) assemblage from two localities of the Argentine Eastern Puna (Calahoyo and Casira) is described. The newly recorded taxa—Cingulata, Dasypodidae, Eutatini: Stenotatus sp. indet. and Eutatini indet., Euphractini: Macrochorobates scalabrinii (Moreno and Mercerat, 1891), and Tardigrada, Mylodontinae: cf. Simomylodon sp. indet. and Simomylodon cf. S. uccasamamensis Saint-André et al., 2010—and those already published from Calahoyo—Cingulata: Macrochorobates chapadmalensis (Ameghino, 1908), Eosclerocalyptus sp. indet., and Tardigrada, Megatheriidae: Pyramiodontherium bergi (Moreno and Mercerat, 1891)—suggest a middle–late Miocene age for the fossil-bearing levels. In Calahoyo, the presence of Stenotatus sp. indet., in addition to some rodents currently under study in the lower levels, suggest a closer similarity with the palaeofauna of Cerdas (southern Bolivia), probably involving the last part of the Miocene Climatic Optimum. The Xenarthra recorded in the middle and upper levels of Calahoyo and Casira suggest a late Miocene–Pliocene age. A comparative analysis between Calahoyo and Casira highlights the absence of Cingulata in the latter and a high diversity in the former. This situation probably indicates different paleoenvironmental conditions. Finally, we present the first certain record of the genus Simomylodon Saint-André et al., 2010 in Argentina, which includes the oldest record of dermal ossicles for sloths in South America.

Tang et al. (2019) described new specimens of carbonaceous compression fossils from the early Cambrian Hetang Formation in South China, for which they established the new taxon Cambrowania ovata Tang and Xiao in Tang et al., 2019. Tang et al. (2019) interpreted these fossils as the remains of metazoans, representing either the carapaces of bivalve arthropods, or early life-cycle stages of sponges. We contest the animal affinity of these fossils; instead, we propose that the specimens described as Cambrowania ovata are actually large Leiosphaeridia—in other words, collapsed hollow organic spheroidal acritarchs. The features described by Tang et al. (2019) all fall into the morphology of carbonaceous compressions of Leiosphaeridia with pyritized/baritized folds and compaction wrinkles. Such Leiosphaeridia are a common component of Cambrian (and older) siliciclastic deposits, and frequently exhibit such a pattern of pyritization, baritization, and encrustation with other diagenetic minerals.

We recently reported Cambrowania ovata Tang and Xiao in Tang et al., 2019, from the early Cambrian Hetang Formation in South China and interpreted it as a problematic animal fossil, possibly related to either sponges or bivalved arthropods (Tang et al., 2019). Slater and Budd (2019) contested our taxonomic identification and phylogenetic interpretation; instead, they claimed that Cambrowania ovata is a large acritarch referable to morphotaxon Leiosphaeridia Eisenack, 1958, and thus is not an animal. Here we refute their criticisms, clarify the differences between Cambrowania and Leiosphaeridia and other acritarchs, and reiterate why an animal affinity for Cambrowania cannot be ruled out.