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

Lagenochitina esthonica is a globally distributed chitinozoan in Early to Middle Ordovician rocks. It is regarded as an index species for the early Floian in North America and has a stratigraphically constrained range in other regions. Lagenochitina esthonica is distinguished from other chitinozoans by a distinct flexure, a nearly rounded-square chamber, and a cylindrical neck with a flaring collar. However, since the first description of the species in the 1950s, it has included two varieties: a relatively short form with a test length ~400 μm, and a slender form usually longer than 600 μm. In order to revise the taxonomy of the L. esthonica group, we carried out a statistical morphometric study of a large collection of well-preserved specimens from the Baltic region where the taxon was first established. Additionally, the stratigraphic and geographic distribution of both forms was analyzed based on available occurrence data. The results show that the short form occurs in the upper Tremadocian to lower Dapingian, whereas the slender form is mostly reported from the lower and middle Darriwilian. Both forms are identified on Baltica; the short form has also been reported from Laurentia and South China, whereas the other is known also from Avalonia and Gondwana. The morphological distinction, together with differences in stratigraphic and spatial ranges, suggest that the two forms represent separate species: the original stout L. esthonica, based on the morphology of the holotype, and the slender L. megaesthonica n. sp., described herein. The updated taxonomy enhances the stratigraphic and biogeographic usefulness of lagenochitinids globally.

UUID: http://zoobank.org/ec49166e-2a8d-4941-8723-f023853c5a7e.

Clathrodictyids are the most abundant stromatoporoids in the Upper Ordovician Xiazhen Formation (middle to upper Katian) of South China. A total of nine species belonging to four clathrodictyid genera are identified in the formation, including Clathrodictyon idense Webby and Banks, 1976, Clathrodictyon cf. Cl. microundulatum Nestor, 1964, Clathrodictyon cf. Cl. mammillatum (Schmidt, 1858), Clathrodictyon megalamellatum Jeon n. sp., Clathrodictyon plicatum Webby and Banks, 1976, Ecclimadictyon nestori Webby, 1969, Ecclimadictyon undatum Webby and Banks, 1976, Camptodictyon amzassensis (Khalfina, 1960), and Labyrinthodictyon cascum (Webby and Morris, 1976). The clathrodictyid fauna in the Xiazhen Formation is very similar to those from both New South Wales and Tasmania, although the latter two Australian regions do not share any common clathrodictyid species during the Late Ordovician. The paleobiogeographic pattern indicates that the northward drift of South China resulted in a favorable environment for the migration of clathrodictyids from other peri-Gondwanan terranes to South China. In addition, these peri-Gondwanan clathrodictyid species hosted various endobionts, representing a variety of paleoecological interactions. The high abundance and species-level diversity of clathrodictyid species presumably increased the substrate availability of suitable host taxa, judging from the diverse intergrowth associations between clathrodictyids and other benthic organisms. These paleoecological interactions between stromatoporoid and other organisms are known from the Late Ordovician and became more abundant and widespread in the Siluro–Devonian. Overall, the Late Ordovician clathrodictyid assemblage in South China demonstrates one of the highest species-level diversities among all peri-Gondwanan terranes and represents a precursor of the complex, clathrodictyid-dominated communities of later metazoan reefs during the Great Ordovician Biodiversification Event.

UUID: http://zoobank.org/6063c47d-cb77-4a03-98cf-b2354656dea6

Cap-shaped skeletal fossils are the earliest undisputed body fossils of mollusks appearing in the basal Cambrian. A study on the morphometry of cap-shaped fossils from the Nanjiang area (North Sichuan, China) is undertaken to understand the origin and evolution of the early mollusks. The distribution of these fossil cap-shaped mollusks indicates a stepwise increase in their diversity during the early Cambrian. Maikhanella Zhegallo in Voronin et al., 1982 co-occuring with the spinose sclerites of siphogonuchitids, is regarded as the earliest scleritized mollusk. It is followed by other maikhanellids, e.g., Purella Missarzhevsky, 1974 and Yunnanopleura Yu, 1987, which co-occur with the earliest univalved helcionellids, e.g., Igorella Missarzhevsky in Rozanov et al., 1969. Cluster analysis of their morphometric characteristics shows that the Maikhanella group is similar to the Purella and Yunnanopleura groups, but is less comparable with univalved helcionellids. The maikhanellids are interpreted as representatives of the stem group Aculifera, although it remains uncertain if one or two larger cap-shaped shell plates were present on the elongate slug-like body, comparable to those of Halkieria Poulsen, 1967 or Orthrozanclus Conway Morris and Caron, 2007. Maikhanellids are characterized by the prominent protrusions or scales on the cap-shaped shell plates arranged in a concentric pattern around the shell apex. Evolutionarily, the protrusions or scales are reduced in younger strata, whereas subsequently a typically concentric ornament developed, the cap-shaped shell plates developed higher profiles, and the apical region became increasingly bare of scales. Meanwhile, the cap-shaped shell plates gradually evolved into a helcionellid-like appearance with an anteroposteriorly inclined apex. The morphological evolution of the earliest sclerotized mollusks reflects biotic evolution and environmental adaption among the stem-group mollusks during the early Cambrian.

A non-biomineralized arthropod, Protocaris marshi, was described from the lower Cambrian (Dyeran Series 2, Stage 4) of Parker's Cobble in northwestern Vermont in 1884. It represents the first fossil exhibiting Burgess Shale-type preservation to have been discovered. The locality was presumed to have been worked out and was not collected in a significant way for more than 100 years. Rediscovery of productive layers has yielded soft-bodied and lightly sclerotized taxa new to the locality, including the alga Fuxianospora, a possible priapulid, a radiodont, and a specimen tentatively assigned to Herpetogaster. New specimens of the sponge Leptomitus zitteli, the bivalved arthropod Tuzoia, and the chordate Emmonsaspis cambrensis provide additional information on those taxa, and multiple specimens allow a bivalved arthropod, Vermontcaris montcalmi new genus, new species, to be described. The primary mode of fossil preservation is as carbonaceous compressions. The Parker Quarry Lagerstätte complements the Kinzers Formation of Pennsylvania (also Series 2, Stage 4) in revealing the diversity of soft-bodied taxa on the southern margin of the paleocontinent Laurentia.

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The fossil record of octocorals from Cenozoic marine strata of western North America is quite limited, and they have not been reported previously from rocks in Washington State, USA. Two late Oligocene specimens from the upper part of the Lincoln Creek Formation in western Washington, referred to Radicipes? sp., are the first fossil record of the family Chrysogorgiidae. The family Isididae is represented by an internode and two holdfasts identified as Isidella sp. collected from the Oligocene Pysht Formation, along with specimens questionably identified as Lepidisis sp., possibly the first fossil record for this genus. Together, these are the first confirmed fossils of the Alcyonacea from north of California in western North America. The axes of sea pens from several late Eocene or early Oligocene localities in the Lincoln Creek Formation in the central part of western Washington, and the Pysht and Makah formations on the north side of the Olympic Peninsula, are the first fossil record for the Pennatulacea from western North America; all are tentatively referred to the genus ‘Graphularia’. Large axes from the Lincoln Creek Formation and Makah Formation are referred to ‘Graphularia’ (?) aff. sasai, because they are similar to the species known only from late Eocene and early Oligocene rocks in Japan.

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).

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

In the early Cambrian fossil record, triradial symmetry is typical for anabaritids and occurs among carinachitids. The former are an extinct group of minute benthic cnidarians covered with a calcareous tubular exoskeleton. The origin of the anabaritids is poorly understood, but previously reported triradial pyramid-shaped steinkerns and molds of the oldest conulariids, Vendoconularia, from the upper Ediacaran of the White Sea region suggested the anabaritids were closely related to conulariids. However, triradial symmetry could originate independently in different lineages in the late Ediacaran and early Cambrian. Herein we describe a new taxon, Ilankirus kessyusensis new genus new species, from the base of the Cambrian Stage 2 of the Olenek Uplift (Siberian Platform). These fossils occur as ornamented steinkerns in the shape of trilateral pyramids and lack any relics of a mineralized exoskeleton. Abundant plastic deformations and fractures of the casts suggest the organism was weakly if at all mineralized. The steinkerns are encrusted with a thin patina of iron-rich chlorite (chamosite) formed because of a multistage diagenetic replacement of authigenic glauconite (glauconite–berthierine–chamosite) under reducing conditions of oxygen-depauperate pore- and seawater. Both lacking two major autapomorphies of the crown-group conulariids (mineralized periderm and quadrate cross section of the oral region of the periderm), the late Ediacaran triradial Vendoconularia and Terreneuvian Ilankirus represent stem-group conulariids.

UUID: http://zoobank.org/a2ce04fa-36a5-485a-806c-f13f4749fc7f

The fossil record of treeshrews, hedgehogs, and other micromammals from the Lower Siwaliks of India is sparse. Here, we report on a new genus and species of fossil treeshrew, specimens of the hedgehog Galerix, and other micromammals from the middle Miocene (Lower Siwalik) deposits surrounding Ramnagar (Udhampur District, Jammu and Kashmir), at a fossil locality known as Dehari. The treeshrew from Dehari (Sivatupaia ramnagarensis n. gen. n. sp.) currently represents the oldest record of fossil tupaiids in the Siwaliks, extending their time range by ca. 2.5–4.0 Myr in the region. Dietary analyses suggest that the new tupaiid was likely adapted for a less mechanically challenging or more frugivorous diet compared to other extant and fossil tupaiids. The occurrence of Galerix has only been recently documented from the Indian Siwaliks and the Dehari specimens help establish the likely presence of a relatively large Siwalik Galerix species in the Ramnagar region. In addition to the new treeshrew and hedgehogs, new specimens of the rodents Kanisamys indicus, Sayimys sivalensis, and Murinae indet. from Dehari help confirm that age estimates for the Ramnagar region are equivalent to the Chinji Formation in Pakistan, most likely corresponding to the middle to upper part of the Chinji Formation.

UUID: http://zoobank.org/56fb160c-2df8-4cd3-be91-af4dc02d0979

The Lower–Middle Ordovician (Ibexian, Whiterockian) sedimentary rocks exposed at Rancho Las Norias includes the informally named Las Norias formation, which consists of an intercalation of carbonate and clastic sediments with abundant marine fauna. These deposits occur as the most austral sedimentary rocks of Ordovician age for Laurentia, providing a critical link to understand the distribution of Ordovician marine faunas of North America. An investigation of the gastropod fauna from the upper portion of the Las Norias formation, Sonora, Mexico, is undertaken for the first time. The gastropod assemblage includes Maclurites acuminatus, ?Monitorella sp., Lecanospira sp., Malayaspira aff. M. rugosa, Lophospira perangulata, and Hormotoma? sp. This assemblage indicates a paleogeographic relationship with Laurentia, including the USA (Nevada), Canada (British Columbia, Newfoundland), Greenland, and the Argentine Precordillera.

The Cambrian stratigraphic succession of the Pyrenees (SW Europe) has undergone a complex Variscan and Alpine tectonothermal history leading to marked metamorphism and development of cleavage networks, which might partly explain the lack of Cambrian fossiliferous beds. This gap has traditionally precluded its paleobiogeographic and biostratigraphic relationships with other neighboring peri-Gondwanan units. Correlations are only based on lithostratigraphic comparisons and radiometric constraints. In this general scheme, the Terrades inlier (Gerona Province, Spain) provides the only significant and indisputable ‘early Cambrian’ fossil record of the Pyrenees. This predominantly siliciclastic outcrop consists of multiple patch reefs and bioherms having yielded archeocyaths dated at Cambrian Epoch 2, Age 3. This paper describes, for the first time, the microfossil assemblage included in the archeocyathan-microbial reefal complex that crops out in the Terrades inlier to clarify its age and affinities with surrounding tectonostratigraphic units. Reefal flanks of patch reefs have yielded bradoriids, brachiopods, molluscs, tommotiids, chancelloriids, hyoliths, and the problematic fossil (and chronostratigraphically significant) Rhombocorniculum cancellatum Cobbold, 1921. In addition to confirming the previously assigned age of the succession, the recovered fauna emphasizes strong affinities with the surrounding Occitan Domain (Montagne Noire, southern Massif Central, France) and Sardinia (Italy). Along with lithostratigraphic comparison and tectonic considerations, this further supports the recent reconstructions positioning the Pyrenean domain between the Montagne Noire (to the southwest) and Sardinia (further to the northeast) on the Gondwana margin during Cambrian times.

Silty noduliferous shales in the lower part of the Middle Ordovician (late Darriwilian 2) Taddrist Formation in the central Moroccan Anti-Atlas contain a unique assemblage of conulariids consisting of one species each of Archaeoconularia Bouček, 1939, Glyptoconularia Sinclair, 1952, and Pseudoconularia Bouček, 1939. Glyptoconularia antiatlasica new species, currently represented by a single three-dimensional specimen, is the first member of this extremely rare, highly autapomorphic genus to be described from outside of cratonic North America as well as the first Glyptoconularia from the Middle Ordovician. Pseudoconularia cf. P. grandissima (Barrande, 1867) and Archaeoconularia cf. A. exquisita (Barrande, 1867) most closely resemble species previously described from Middle and Upper Ordovician strata in the Prague Basin (Czech Republic), then located adjacent to North Africa. Anaconularia anomala (Barrande, 1867), previously known from the Upper Ordovician (Sandbian) Libeň and Letná formations in the Prague Basin, is documented for the first time from the Upper Ordovician (Katian) Upper Tioririne Formation of Morocco. In addition to extending the known stratigraphical and paleogeographical ranges of these conulariids, results of the present investigation add to the list of invertebrate taxa that appear to have originated during Early or Mid-Ordovician times in peri-Gondwana or south-polar Gondwana, and then migrated to the eastern margin of Laurentia, arriving there along with other cool-water taxa during the Sandbian-Katian transition.

UUID: http://zoobank.org/9ea5051b-cff5-4911-861a-531db4f45883

Strata of the Solvik Formation in the central Oslo region (upper Hirnantian to upper Aeronian) are rich in atrypides that elucidate the evolution of this group. A few athyridide brachiopods are also present. Eleven taxa of smooth spire-bearers are described taxonomically using fossils and peels. Among these occur one new genus, Eisaella, and five new species: Eisaella uniplicata, Thulatrypa huangi, T. vikenensis, ?Becscia pentagona, and Shelvothyris bivittata. Atrypides make up 30% (24 genera) of the total brachiopod genera in the Solvik Formation, more than one third of the known global atrypide fauna during that interval. Most are members of the family Lissatrypidae, illustrating the evolution of this group from upper Hirnantian and lower Silurian strata. These show close affinity to each other attesting to ongoing radiation. The new genus Eisaella is a likely ancestor of Lissatrypa, while Meifodia displays a gradual evolution through the formation. Atrypides were most diverse at the base and top of the Solvik Formation due to heterogony in environments and water depth, but were less diverse in the comparatively constant environmental setting of the late Rhuddanian time. The basal assemblages with few atrypides from shallow facies may be compared to the shallow Cathay Fauna of China. Most of the abundant Norwegian atrypide fauna is found in deeper water with few comparable faunas known globally. The Oslo region may have served as a center for the evolution and spread of atrypides immediately following the end-Ordovician glacial drawdown in sea level.

UUID: http://zoobank.org/508dba95-5501-403d-9a4d-deeb324847c9

The diversity of mid-Cretaceous tyrannosauroids is poorly understood. We describe a partial tyrannosauroid femur from the Albian–Cenomanian Wayan Formation of eastern Idaho that helps to fill in an important spatiotemporal gap in the North American record of tyrannosaurs. This specimen, consisting of the proximal half of the bone, is morphologically similar to the femur of Moros intrepidus, a small-bodied tyrannosauroid from the Cenomanian Mussentuchit Member of the Cedar Mountain Formation of Utah, but not referable to this taxon. The Wayan femur lacks an autapomorphy diagnostic for Moros intrepidus, indicating the presence of a previously unrecognized tyrannosauroid taxon in the early Late Cretaceous of Laramidia. Histological results indicate that, at the time of death, this individual was at least five years old, skeletally immature, and undergoing growth at a moderate rate. The addition of this tyrannosauroid to the Wayan-Vaughn Assemblage provides additional evidence for the widespread distribution of various tyrannosauroid taxa in Laramidia during the early Late Cretaceous.

An assemblage of 34 chiton valves collected from rocks of the “Imperial” Formation sensu lato at Super Creek near Palm Springs, California, is described here. The sedimentary rocks exposed at Super Creek were deposited in the proto-Gulf of California during the Late Miocene. This chiton assemblage represents the first reported Miocene chitons from western North America. The chiton valves are classified as Callistochiton cf. C. elenensis, Chaetopleura cf. C. lanuginosa mixta, Calloplax roederi n. sp., Chiton solaris n. sp., Callistoplacidae sp. indet., and Chaetopleuridae sp. indet. Thick valves dominate the assemblage. The chiton fossils reflect the overall faunal pattern from the “Imperial” Formation s.l. of a strong similarity to taxa in the modern Gulf of California, with a Caribbean component as well. This assemblage fills a gap in the chiton fossil record and preserves details of the adaptive radiation of the Polyplacophora that occurred at the time in concert with the onset of an upwelling regime.

UUID: http://zoobank.org/9947bfc9-84a3-4358-bac7-e660b936b067

Herein is reported the first occurrence of ostracodes from the Iapó Formation, an uppermost Ordovician unit of the Rio Ivaí Group in the Paraná basin, Brazil. Two ostracode species were identified in the Três Barras Farm section: Harpabollia harparum (Troedsson, 1918) and Satiellina paranaensis Adôrno and Salas in Adôrno et al., 2016 were recovered from dropstone-bearing shale overlying glaciogenic diamictites, a feature typical of Hirnantian (uppermost Ordovician) strata throughout Gondwana. The taxonomy of the Genus Harpabollia, as well as its type species Harpabollia harparum, was reviewed, and emended and new diagnoses were respectively proposed for each taxon. Occurrences of Harpabollia harparum and Satiellina species were common in areas influenced by cold waters. Additionally, the occurrence of Harpabollia harparum, an index species to the uppermost Ordovician of several stratigraphic units in Baltica and southern Gondwana, allowed us to infer a Hirnantian age for the deposits of the Iapó Formation. Other than being associated with Harpabollia harparum in Iapó Formation of the Paraná basin, Satiellina paranaensis is also found in lower levels of the Vila Maria Formation; therefore, these are also considered Hirnantian in age. Above these lower levels of the Vila Maria Formation, a well-dated Rhuddanian (lowermost Llandovery, Silurian) palynomorph assemblage is observed within the formation. These occurrences are evidence of a continuous process of sedimentary deposition during the Ordovician–Silurian transition in the Paraná basin.

We report the finding of two partial specimens of Cryptolepas rhachianecti (Cirripedia, Coronulidae), a coronulid barnacle known only to inhabit the skin of gray whales (Eschrichtius robustus), in Pleistocene-aged sediments from the Canoa Basin, Ecuador. While the historical range of gray whales includes the North Pacific and North Atlantic, to our knowledge this is the first inferred evidence of a gray whale population having resided within the South Pacific. We describe the two Cryptolepas rhachianecti fossils, use isotopic analysis to investigate evidence of migration in their host whales, and discuss their implications for our understanding of gray whale evolutionary history.

A new genus and species of microconchid tubeworm, Aculeiconchus sandbergi n. gen. n. sp., is described from the Givetian (Devonian) Maywood Formation of Cottonwood Canyon, Wyoming, USA. It possesses unique hollow spines of various lengths on the tube underside, a position previously undocumented for these fossils. Like some cyclostome bryozoans possessing basal tubular extensions, the basal spines of Aculeiconchus n. gen. were presumably also used for fixation to flexible substrata, e.g., algal thalli, which is a previously undocumented adaptive strategy in microconchids. Together with other skeletal features, such basal spines could suggest that ‘lophophorate’ microconchids, unlike the other tentaculitoids, might be phylogenetically not as distant from bryozoans as previously thought. The Maywood Formation, which contains a few-millimeters thick, monospecific shell accumulation of the microconchids described herein, records deposition in an estuarine brackish setting within narrow channels that were cut into underlying strata. The microconchids were opportunistic taxa that repeatedly colonized these salinity-stressed estuarine channels, leading to a series of adaptive innovations, including colonization of plant stems during the Early Devonian (Beartooth Butte Formation) and possibly flexible, soft-algal substrata during the Middle Devonian (Maywood Formation, this study). Tectonic quiescence during the Early and Middle Devonian indicates that erosion and subsequent deposition of the Maywood and the underlying Beartooth Butte Formation channels were responses to major eustatic events. Over a span of nearly 30 Myr, channels were cut successively during lowstand conditions and a distinctive faunal assemblage with microconchids tracked marine transgressions into the channels.

UUID: http://zoobank.org/394c8b32-d5e7-411e-8e56-6fb9f55bbb8a