A new cornulitid genus and species, Porkuniconchus fragilis new genus and species, is here described from the Ärina Formation (Hirnantian, Porkuni Regional Stage) of northern Estonia. This new taxon differs from most cornulitids by having a fusiform ornamentation pattern that is somewhat similar to that of Kolihaia. All studied specimens are attached to a carbonate hardground. The hardground fauna is by abundance and encrustation area dominated by cornulitids. Other encrusters are represented only by a single sheet-like cystoporate bryozoan. The cornulitid specimens represent different growth stages, which suggest that the hardground was continuously colonized by cornulitid larvae. The high encrustation density indicates that the studied hardground may have represented a high-productivity site in the Hirnantian of the Baltic Basin.

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Predation is a behavior that is commonly unsuccessful, but the cause of failure is often difficult to determine in the fossil record. Here, we report on gastropod drill holes in two Plio- and Miocene bivalve specimens from the Netherlands created from the inner side of the bivalve prey's shell, which we call reverse drill holes. These holes are unequivocally caused by failure of the gastropod drilling predators to make effective use of their chemoreception and mechanoreception sensory adaptations. We hypothesize that the diffuse nature of chemical cues emanating from dense aggregations of living prey could have confused foraging predators and stimulated them to initiate the drilling process on empty valves. Poor decision making due to hunger is an alternative hypothesis. These traces represent the first reported examples of reverse gastropod drill holes from the fossil record, and the first attributed to Naticidae. Compared to other types of failed predation (incomplete drill holes and drill holes in multiply-drilled specimens) in the two assemblages studied, reverse drill holes are rare (< 1% of drill holes). This result implies that the driller's sensory and decision-making processes were generally reliable at distinguishing dead from live prey.

The Eager Formation (Cambrian Stage 4) of the Cranbrook area, British Columbia, contains abundant, sometimes beautifully preserved, trilobites and rare non-biomineralized taxa. Trilobites were first reported just over a century ago but have received little research attention, resulting in uncertainty in the number and identity of species within the assemblage and ambiguity in the age of the fauna. The trilobites of the Eager Formation in the Cranbrook area are described herein based largely upon material collected in 2015. A minimum of eight (and perhaps up to 11) trilobite species are recognized. The four species of olenelloids (Olenellus santuccii Webster n. sp., Olenellus? schofieldi, Mesonacis eagerensis, and Wanneria cranbrookense Webster n. sp.) are by far the most abundant elements of the fauna. At least two, and perhaps as many as five, species of dorypygid are present, as are two species of “ptychoparioids”. Paucity and poor preservational quality of specimens mean that the various dorypygid and “ptychoparioid” species are left in open nomenclature. Trilobite diversity in the Cranbrook Lagerstätte is comparable to that within other Lagerstätten from Cambrian Stage 4 (Series 2) of Laurentia. The diversity and abundance of trilobites, combined with biostratinomic and trace fossil data, suggest that the assemblage is autochthonous and/or parautochthonous, and that the local environment was at least periodically able to support a “typical” benthic trilobite community. The age of the Cranbrook Lagerstätte is constrained to lie within the middle Dyeran Stage of Laurentia, within what was a substantial stratigraphic gap in the distribution of Burgess Shale-type Lagerstätten.

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One of the challenges for bryozoans is to avoid refiltering water that has already had its plankton removed. Larger colonies develop colony-wide maculae-centered feeding currents to avoid refiltering water and generally have elevated maculae (monticules). We hypothesize that the height and density of spacing of monticules are inversely proportional to curvature of the colony surface. Larger, flatter colonies should have higher and more closely spaced monticules. We compare two Permian stenolaemate bryozoans whose colonies form branches with elliptical cross sections: the smaller and more elliptical cystoporate Evactinostella crucialis (Hudleston, 1883) from Western Australia (N = 17) and the larger and flatter trepostome Tabulipora sp. from eastern North Greenland (N = 15). Using calipers and digital elevation models, we measured curvature, monticule height, and number of monticules per area. Results indicate that Evactinostella branches are at least twice as curved as those of Tabulipora, their monticules are half the height of Tabulipora, and their monticules are 22% less densely spaced than those of Tabulipora. In Evactinostella colonies, surface curvature is inversely proportional to monticule height and spatial density, which is not true for Tabulipora. Therefore, we conclude that the smaller and more curved the colony surface, the less the colony needs robust colony-wide feeding currents created by tall, closely spaced monticules.

Dysnoetoporidae is a family of cheilostome bryozoans including only the genus Dysnoetopora Canu and Bassler, 1926, with three recognized species from the Late Cretaceous of the United States, Crimea, and Germany. The aim of this study was to record for the first time the presence of a fossil dysnoetoporid bryozoan in South America by describing Chenquepora miocenica new genus new species. This taxon was found in Miocene deposits of the Chenque Formation (early Langhian, ~15.37 Ma), on the Atlantic coast of Argentine Patagonia. Chenquepora n. gen. differs from Dysnoetopora in its encrusting colony and, consequently, in the absence of an endozone with long zooids arranged parallel to growth direction. This new record extends its paleobiogeographic distribution to the Southern Hemisphere and its stratigraphic range from the Campanian (Late Cretaceous) to the Miocene, showing that this extinct family survived the Cretaceous-Paleogene (K-Pg) mass-extinction event.

The early to middle Permian brachiopods from the Tengchong Block in western Yunnan, southwestern China, play important roles in biostratigraphic correlation and paleogeographic inferences of tectonic units on eastern peri-Gondwana. However, detailed taxonomic studies of these brachiopods have been limited. In this paper, we provide the systematic description of three Permian brachiopod assemblages from the Shanmutang section in the northern Tengchong Block, which in ascending order include the Elivina-Etherilosia Assemblage from the top of the Kongshuhe Formation, and the Spiriferella-Spiriferellina Assemblage and the Waagenites-Costiferina Assemblage from the base and lower part, respectively, of the overlying Dadongchang Formation. Based on the biostratigraphic assessment of the brachiopod taxa as well as the age constraints from the associated fusulinid assemblages, the age of the Elivina-Etherilosia Assemblage is considered to be most likely late Sakmarian to early Artinskian, the Spiriferella-Spiriferellina Assemblage probably late Artinskian, and the Waagenites-Costiferina Assemblage late Roadian to early Wordian. A transition from cool-water faunas with Gondwanan affinities to mixed warm-temperate (transitional) faunas with Cathaysian elements is evident for the Artinskian and through the Guadalupian. The climatic transition inferred from the faunal evidence is interpreted to have resulted from the combined effect of Gondwana deglaciation and the northward drift of the peri-Gondwanan blocks (including the Tengchong Block) during the Cisuralian and Guadalupian.

The ontogeny of two species of corynexochid trilobites from the middle Cambrian Mount Cap Formation of the eastern Mackenzie Mountains, northern Canada, is documented. Sahtuia carcajouensis (Dolichometopidae) and Mackenzieaspis parallelispinosa (Zacanthoididae) are both endemic to this formation and only known from one locality. They, along with several other corynexochid taxa, occur in a succession of mudstone with scattered carbonate interbeds, deposited in a weakly storm-agitated setting near the flank of a semi-enclosed basin. The ontogeny of both species is characterized by mainly normal cranidial development, but a unique distribution of segments in their thoraxes and pygidia. The number of trunk segments was typical for their respective families, whereas the final number of segments released into the thorax was reduced. This occurred in both species through timing modifications to segment release, indicating heterochrony. Sahtuia carcajouensis and Mackenzieaspis parallelispinosa are likely derived from two separate clades, and heterochrony probably arose separately but synchronously. The endemicity of both species probably reflects unique paleoecological conditions in this part of the basin. Preliminary results indicate that the fossil-bearing mudstone was deposited under well-oxygenated conditions that underwent high nutrient flux and possibly experienced varying salinity. These factors may have affected the organisms’ physiology, or perhaps provoked an adaptation to achieve early maturation.

The terminal Ediacaran Shibantan biota (~550–543 Ma) from the Dengying Formation in the Yangtze Gorges area of South China represents one of the rare examples of carbonate-hosted Ediacara-type macrofossil assemblages. In addition to the numerically dominant taxa—the non-biomineralizing tubular fossil Wutubus and discoidal fossils Aspidella and Hiemalora, the Shibantan biota also bears a moderate diversity of frondose fossils, including Pteridinium, Rangea, Arborea, and Charnia. In this paper, we report two species of the rangeomorph genus Charnia, including the type species Charnia masoni Ford, 1958 emend. and Charnia gracilis new species, from the Shibantan biota. Most of the Shibantan Charnia specimens preserve only the petalodium, with a few bearing the holdfast and stem. Despite overall architectural similarities to other Charnia species, the Shibantan specimens of Charnia gracilis n. sp. are distinct in their relatively straight, slender, and more acutely angled first-order branches. They also show evidence that may support a two-stage growth model and a epibenthic sessile lifestyle. Charnia fossils described herein represent one of the youngest occurrences of this genus and extend its paleogeographic and stratigraphic distributions. Our discovery also highlights the notable diversity of the Shibantan biota, which contains examples of a wide range of Ediacaran morphogroups.

UUID: http://zoobank.org/837216cd-4a4a-4e13-89e2-ee354ba48a4c

The Cambrian dokimocephalid trilobite genus Deckera Frederickson, 1949 is a minor component of Steptoean (Jiangshanian) faunas of Laurentian North America. The original diagnosis emphasized strongly inflated palpebral areas of fixed cheeks and elevated palpebral lobes as important and novel characters. An appraisal of archival and new specimens from Oklahoma, Missouri, Pennsylvania, Nevada, Utah, and Newfoundland show that the genus is geographically widespread and likely represented by as many as nine species, although only two of them are named formally. Deckera cf. D. aldenensis Frederickson, 1949 from Nevada extends the stratigraphic range of Deckera down to the base of the Jiangshanian Stage, and new species from Nevada and Newfoundland take the genus down farther, into upper Paibian strata. Paibian species show that some basal members of the genus have weakly inflated, nearly flat palpebral areas with palpebral lobes that sit well below the crest of the glabella. A revised diagnosis of Deckera focuses on the broad cranidium with width across the palpebral lobes much greater than the sagittal length. Pygidia are known for a few species and all of them have an unusual flexure of the posterior margin and border. Lectotype and paralectotype specimens are designated for Deckera completa.

The existence of Ordovician Peruvian cephalopods has been known since at least the 1910s. However, they have not been effectively documented previously with only a few described taxa listed in open nomenclature. Here, we describe a cephalopod assemblage at the finest taxonomic level possible. The specimens were collected from the Floian section (Baltograptus minutus graptolite Zone) of the San José Formation from the Kimbiri area, northwest of Cuzco (= Cusco), and from a section along the Inambari River, southeastern Peru. The dark mudstone-siltstone of the San José Formation was deposited within the Central Andean Basin. The assemblage contains five species of small orthoceracones belonging to four families and three orders, consisting of one indeterminate dissidocerid, one bathmoceratid (Saloceras sp.), one rioceratid (Rioceras? sp.), and two baltoceratids belonging to Annbactroceras grecicostatum (Kobayashi, 1937), and Bactroceras cocafolium new species. The dominance of small orthoceracones is typical for early Paleozoic pelagic cephalopod assemblages. One species, A. grecicostatum, is known from elsewhere in the Central Andean Basin. The other taxa indicate a peri-Gondwana-Avalonia paleogeographical relationship of the cephalopod fauna, which is consistent with previously published data from brachiopods and trilobites.

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The trilobite faunas that occur with the Steptoean Positive Isotope Carbon Excursion (SPICE) at Smithfield Canyon, Utah, have been reported, but not illustrated. Given the importance of the SPICE at this section for international correlations, the trilobites from new collections from the upper Nounan Dolomite to lower St. Charles Formation at Smithfield Canyon are reported herein and integrated with the previously reported taxa. Trilobite assemblages indicate that the upper Cedaria to the Ellipsocephaloides biozones (Miaolingian Series, Guzhangian Stage to Furongian Series, Jiangshanian Stage) are present stratigraphically below or above the SPICE.

Some of the taxa reported herein may represent new species, but they are not represented by well-enough preserved specimens and are left in open nomenclature. However, Kingstonia smithfieldensis n. sp. and Bromella utahensis n. sp. are named on the basis of common and well-preserved specimens.

New carbon isotope data from Smithfield Canyon from an overlapping section of the lower St. Charles Formation, that add to the overall shape of the SPICE curve, are presented. The new δ13C values above the Elvinia Biozone range from –0.36‰ to +1.5‰, confirming that the SPICE concludes within the Elvinia Biozone.

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Decimeter-scale, elongate, fossil fronds from the Ediacara Range in South Australia were formally described as Rangea longa Glaessner and Wade, 1966, but the disparate nature of documented specimens has hindered their inclusion in global syntheses and has resulted in these fossils being assigned to at least five different genera in two different clades since their discovery. Detailed study of the type material from the Ediacara Range and the few specimens subsequently collected elsewhere in the Flinders Ranges reaffirms that these specimens represent a single species, with the apparent morphological variation between specimens entirely taphonomic and reflecting the obverse and reverse surfaces of these fronds coupled with the orientation of the frond axis and petaloids at different angles relative to the sea bottom on which they were preserved. The preserved architecture of these fronds constitutes three orders of branching microstructure that are strictly orthogonal to immediately higher and lower orders. This implies affinities with the arboreomorphs, but representing a new frond genus herein named Akrophyllas. Akrophyllas n. gen. differs from all other Ediacaran fronds in exhibiting a stalk that is visible only on one side of the frond and is internal to the other side where the first-order branches instead meet at a zigzag axial trace. Akrophyllas n. gen. was attached to a bulbous holdfast on the sea bottom, and evidence for current scours that formed in the lee of the fronds and for a strong current alignment of felled fronds with depositional overlap of adjacent fronds imply an upright, epibenthic lifestyle for Akrophyllas longa new combination.

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A new genus and species of arachnid (Chelicerata: Arachnida), Douglassarachne acanthopoda n. gen. n. sp., is described from the late Carboniferous (Moscovian) Coal Measures of the Mazon Creek Lagerstätte, Illinois, USA. This is a unique animal with distinctive large spines on the legs. It has a subovate body, a segmented opisthosoma, and a terminal anal tubercle. The legs are robust and appear to have been similar in construction throughout the limb series, with heavy spination of the preserved proximal podomeres. The mouthparts and coxo-sternal region are equivocal. The preserved character combination does not permit easy referral to any known arachnid order, living or extinct, thus the new fossil in placed as Arachnida/Pantetrapulmonata incertae sedis. It contributes to an emerging pattern of disparate body plans among late Carboniferous arachnids, ranging from anatomically modern members of living orders through to extinct taxa, such as the present fossil, whose phylogenetic position remains unresolved.

UUID: http://zoobank.org/b70f5f95-9c8b-4389-bee5-b6031bff2ee2

Natural history museums house numerous previously undescribed species and unknown information hidden in their collections. We describe lower Carboniferous slit-bearing gastropods (order Pleurotomariida, subclass Vetigastropoda; and family Goniasmatidae, subclass Caenogastropoda) from previously unreported gastropod collections made by Jane Longstaff (Jane Donald), one of the pioneering paleontologists of Paleozoic gastropods in the late nineteenth and early twentieth centuries. The gastropods were collected from the Lower Limestone Formation (Visean, Brigantian) near Dalry, Ayrshire, Scotland. The collection consists largely of microgastropods, many of which are unusually well-preserved including delicate ornament and protoconchs (larval shells). Three new pleurotomariidan species are described—Biarmeaspira heidelbergerae new species, Neilsonia seussae new species, Tapinotomaria longstaffae new species—in addition to seven species belonging to Borestus Thomas, 1940, Stegocoelia (Stegocoelia) Donald, 1889, Stegocoelia (Hypergonia) Donald, 1892, Donaldospira Batten, 1966, and Platyzona Knight, 1945. The caenogastropod-type protoconch is documented for the first time in Hypergonia, which is therefore placed in Goniasmatidae. The new data confirm that Neilsonia Thomas, 1940 (type genus of Neilsoniinae) belongs to Pleurotomariida and is distinct from the morphologically convergent Peruvispira Chronic, 1949 (Goniasmatidae). The selenizone morphology is identical in Biarmeaspira Mazaev, 2006 and Baylea de Koninck, 1883 during their early ontogeny, and Biarmeaspira develops an angulation on the selenizone (the diagnostic feature) in late ontogeny. This corroborates earlier suggestions that Biarmeaspira evolved from Baylea. Biarmeaspira heidelbergerae n. sp. is the first Carboniferous record of Biarmeaspira, which was previously only known from the Permian. The angulated selenizone evidently evolved several times in Pleurotomariida and the repeated appearance of this character in different groups (e.g., Phymatopleuridae, Eotomariidae, Pleurotomariidae) needs further studies using phylogenetic methods.

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Pterotheca Salter, 1853 is an unusual but readily identifiable bellerophontoid gastropod that occurs in the Upper Ordovician to the Llandovery of the lower Silurian in many parts of North America and Europe. Recently, a large collection of Pterotheca was obtained for the first time from the Xiushan Formation of middle Telychian (Llandovery) age in the Hunan Province of South China. This is also the first record of the genus in the low-latitude peri-Gondwanan region. On the basis of the collection, two new species of Pterotheca—P. yongshunensis and P. xiushanensis—were identified and are described herein. The morphologic analysis suggests that close relatives of these new species may be Pterotheca species from the Telychian of Scotland. The new species show continuous variations of marginal apex to submarginal apex, implying that one of the Pterotheca species may be ancestral to the Devonian Aspidotheca Spriesterbach, 1919. The Pterotheca species from South China possibly lived a slowly crawling life on a soft substrate, feeding on algae and/or detritus, and were adapted to a shallow-water setting with substantial terrigenous input. Given that all the known Silurian Pterotheca species occurred in siliciclastic settings, most of which represent sea-level fall and lowstand periods, we demonstrate that geographic isolation and enhanced ocean circulation during the early Silurian regression facilitated the speciation of Pterotheca globally, and the connection of a sea pathway during the Rhuddanian transgression after the end-Ordovician glaciation could have led to the primary dispersal of Silurian Pterotheca.

Phylogenetic analysis demonstrates that Kuamaia lata, a helmetiid euarthropod from the lower Cambrian (Series 2, Stage 3) Chengjiang Konservat-Lagerstätte, nests robustly within Artiopoda, the euarthropod clade including trilobitomorphs. Microtomography of new specimens of K. lata reveals details of morphology, notably a six-segmented head and raptorial frontal appendages, the latter contrasting with filiform antennae considered to be a diagnostic character of Artiopoda. Phylogenetic analyses demonstrate that a raptorial frontal appendage is a symplesiomorphy for upper stem-group euarthropods, retained across a swathe of tree space, but evolved secondarily in K. lata from an antenna within Artiopoda. The phylogenetic position of K. lata adds support to a six-segmented head being an ancestral state for upper stem- and crown-group euarthropods.

Haliotis Linnaeus, 1758, a commercially important gastropod, is the only known genus in the family Haliotidae (Mollusca, Vetigastropoda, or abalone) worldwide. Its poor Cenozoic record and high intraspecific variability resulted in different interpretations of nomenclature, impeding a robust species-level taxonomy and biogeographic history. Among the best-studied forms, three subspecies of H. tuberculata Linnaeus, 1758 currently inhabit the temperate waters of the Mediterranean and the eastern Atlantic. New findings in the Pliocene of Tuscany (Italy) are presented here, and the taxonomy of the European record is revised. On the basis of a multivariate analysis of shell morphometrics for the first time applied to the study of fossil abalones, and consistent with the chronostratigraphic and geographic framework, H. plioetrusca n. sp. is introduced and H. volhynica Eichwald, 1829 and H. lamellosoides Sacco, 1897 are reinstated as valid species. Some recently described forms from the Pliocene of Spain are placed in synonymy with H. lamellosoides. Haliotis ovata Michelotti, 1847 is proposed as the ancestral taxon of modern H. tuberculata, via H. lamellosoides. This lineage diversified in the subtropical/warm temperate Pliocene Mediterranean, represented by H. lamellosoides, H. bertinii Forli et al., 2003 and H. plioetrusca. The progressive global cooling starting at around 3.0 Ma is associated with the appearance of H. tuberculata at temperate latitudes. H. plioetrusca is not known from younger strata, whereas H. bertinii survived into the Calabrian.

UUID: http://zoobank.org/7c2f2258-2574-4976-a2f5-804c54c86679

We analyzed the ontogenetic trajectories of conch morphology and septal spacing between successive chambers in Cretaceous ammonoids (suborders Perisphinctina and Ancyloceratina) collected from southern India, Madagascar, and Japan. All examined species, except for the family Collignoniceratidae, exhibited similar characteristics during early ontogeny. The common ontogenetic trajectories of septal spacing show a cycle comprising an increase and a subsequent decrease in septal spacing during early ontogeny. The conch diameters at the end of the cycle were estimated to be 1–4 mm. The conch shape (aperture height and whorl expansion rate) covariably changed at this conch diameter. Such covariable changes are commonly recognized in the suborders Perisphinctina and Ancyloceratina. The similarity in the ontogenetic trajectories of conch morphology implies a closer phylogenetic relationship between these suborders compared to Lytoceratina or Phylloceratina.

The terminal Ediacaran Period is signaled worldwide by the first appearance of skeletonizing tubular metazoan fossils, e.g., Cloudina Germs, 1972 and Sinotubulites Chen, Chen, and Qian, 1981. Although recent efforts have focused on evaluating the taxic composition and preservation of such assemblages from the southwestern United States, comparable forms reported in the 1980s from Mexico remain to be re-examined. Here, we reassess the latest Ediacaran skeletal materials from the La Ciénega Formation of the Caborca region in Sonora, Mexico, using a combination of analytical methods: optical microscopy of extracted fossils, thin-section petrography, scanning electron microscopy and energy dispersive X-ray spectroscopy, and X-ray tomographic microscopy. From our examination, we conclude that the La Ciénega hosts a polytaxic assemblage of latest Ediacaran tubular organisms that have been preserved through two taphonomic pathways: coarse silicification and calcareous recrystallization preserving finer details. Further, these fossils show signs that their shells might not have been inflexible or completely mineralized in vivo, and that they might also record tentatively interpreted predation traces in the form of drill holes or puncture marks. This work, along with ongoing efforts around the world, helps to provide a framework for biostratigraphic correlation and possible subdivision of the Ediacaran Period, and further shapes our view of metazoan evolution and ecology in the interval directly preceding the Cambrian explosion.

A new dalmanitid trilobite is described from the early Silurian (Telychian) Waukesha Biota in Wisconsin, USA, best known for its variety of exceptionally preserved arthropods. Waukeshaaspis eatonae new genus new species is the most common trilobite found in the Waukesha Biota, numbering ~200 specimens, allowing a thorough description of the dorsal exoskeleton. This new taxon has a combination of characters unique to the Dalmanitidae, including a distinct embayment on the posterior margin of the pygidium. Embayments are rare for dalmanitids, most species having instead a caudal spine. The lengthening of the genal spines might have served a similar function to that which a caudal spine would have served, allowing the embayment to fulfill a different function, related to keeping an opening when enrolled. The abundance at the preservation site suggests physiological tolerance at the habitation site.

UUID: http://zoobank.org/50a214e3-72f0-4ca2-b8f1-4ad4460bc1d4