Crinoids and blastoids,platyceratid gastropods and time: A taphonomic progression

Taphonomy aims to identify the history of preservation of a fossil, such as the order in which events occurred after death of the organism. For example, why are crinoid and blastoid thecae of Mississippian (Chadian) age never preserved in close association with parasitic/coprophagic platyceratid snails at Salthill Quarry, Clitheroe, Lancashire? They do occur in close juxtaposition at other localities globally where Upper Paleozoic strata crop out. We suggest that the intimate blastoid or crinoid/platyceratid association may occur at one of three identifiable levels. Level 1 includes those rare specimens of crinoids (not known in blastoids so far) that retain all arms and an attached platyceratid, a pattern of preservation indicating rapid burial causing death. Level 2 includes those thecae that have lost their brachioles (blastoids) or arms (crinoids), but still have an attached platyceratid. That is, the echinoderm has started to disarticulate following death, but the platyceratid continues to feed on the theca. In other words, and strangely, the snail is more intimately linked to the theca than certain of the echinoderm's appendages. Level 3 includes sites where crinoid/blastoid thecae and platyceratid conchs are preserved separately; determining if they were previously closely associated may be problematic unless attachment scars of snails are apparent on crinoid thecae. Suggestions that the pelmatozoan/platyceratid association involved aerosol filtration by the snail seem unlikely, because the intimate association is maintained after loss of the arms.     

Palaeozoic micro-pelmatozoan thecae from the bedload of the River Maas (province of Limburg,the Netherlands)

In the Netherlands, Late Palaeozoic pelmatozoans – that is, stalked echinoderms – are known from building stones and cobbles in rivers, but there are no in-situ carbonate rocks from which they might be collected. Unsurprisingly, most recognisable specimens are columnals and pluricolumnals. Two small thecae, collected in the mid-1970s from silexite cobbles in the bedload of the River Maas in the Venlo-Tegelen area (province of Limburg, south-east Netherlands) are exceptional finds. One specimen, the diplobathrid camerate crinoid Rhodocrinites sp., has an unsculptured theca and some minor differences of form, yet otherwise satisfies the diagnosis of this genus. The other, the pentremitid blastoid Doryblastus? sp. is rather poorly preserved, yet is the first blastoid to be recorded from the Netherlands. Either or both of these specimens may be juveniles, particularly the blastoid. They are unlikely to be coeval, coming from separate cobbles and being of slightly different preservation. Their provenance from silexite cobbles suggest they originated from Lower Carboniferous (Tournaisian-Visean = Mississippian) carbonates in the southern Ardennes (south-central Belgium).     

Two intriguing pluricolumnals (Crinoidea) from the Lower Palaeozoic of Powys and Shropshire

Complete fossils must be preferred to fragments for most palaeontological studies, but disarticulated specimens are nonetheless potential sources of noteworthy data. Two crinoid pluricolumnals are recorded from the lower Palaeozoic; informed discussion shows each is a basis for palaeobiological interpretation. Both are gracile and are probably belong to disparids. Floricrinus (col.) sp. is from the Silurian of Wenlock Edge, Shropshire, either from the Much Wenlock Limestone Formation (Wenlock) or, more likely, the Lower Elton Formation (Ludlow). This is the first crinoid from the Silurian of the British Isles with a pentapetaloid arrangement of the areola, a geometry common in the Middle-Upper Ordovician and higher in the geologic column. Pluricolumnal gen. et sp. indet. is from the Lower Llanvirn of Powys. One end of the otherwise straight specimen is tightly coiled. This is likely the proxistele, the most flexible region of the column, and the coiling occurred after the crown was lost by autotomy in response to an environmental disturbance.     

Conodonts of the Mississippian/Pennsylvanian boundary interval in Central Iran

The record of conodonts related to the Mississippian/Pennsylvanian boundary interval was investigated in four sections in Central Iran from two different structural units. Two sections from the Sanandaj–Sirjan trend zone (Asad-abad, and Darchaleh sections) and two from the East-Central Iran Microplate (Shesh-angosht and Kale-Sardar sections) exhibit a nearly complete record previously described across the Mississippian/Pennsylvanian boundary in Iran. The investigated sections can be subdivided in three formations (Ghaleh-, Absheni-, and Zaluda Formation) which belong to the Sardar Group. The mid-Carboniferous boundary was defined by the occurrence of Declinognathus noduliferus s.l.. Bio-event characteristics of the Carboniferous conodont fauna (Mississippian genera Gnathodus and Lochriea have been replaced by Pennsylvanian genera Declinognathus and Idiognathodus) as well as sedimentological changes within overall shallow water deposits were located approximately 33° S of the paleoequator and suggest sea-level changes within the framework of the Late Paleozoic Ice Age (LPIA). Furthermore, a widespread crinoid marker horizon previously described from two localities in Iran can be subdivided into three units of different ages.     

Mississippian foraminifers from the Onimaru Formation in the Hikoroichi area, South Kitakami Belt, Northeast Japan

Abundant foraminifers were found from the Mississippian Onimaru Formation distributed in the Hikoroichi area of the central part of the South Kitakami Belt, Northeast Japan. They include Brunsia pulchra, Archaediscus sp., Paraarchaediscus? sp., Neoarchaediscus? sp., Palaeotextularia spp., Climacammina, spp., Tetrataxis spp., Haplophragmella sp., Lituotubella? sp., Forschiella sp., Cribrospira sp., Bradyina spp., Janischewskina sp., Endothyra spp., Planoendothyra aljutovica., Endothyranopsis compressa, E. crassa, Omphalotis samarica, Eo-staffella spp., Mediocris breviscula, and several others. The Onimaru foraminiferal fauna is similar to those re-ported from MFZ (Mamet Foraminiferal Zone) 15-16. This supports a late Visean (V3b-V3c) age of the formation, which has been proposed previously by rugose corals.     

Notes on Mississippian echinoderms from Hurdlow,Derbyshire, central England

The fossil echinoderms of the Lower Carboniferous (Mississippian) of Derbyshire remain understudied, principally due to the nature of the preservation rather than any lack of biodiversity. Echinoids and crinoids are described in float blocks of limestone from near Hurdlow, Derbyshire, which have been etched naturally after many years of being washed by weakly acidic rain. Surface detail is variable, commonly poor, but rare specimens retain enough features for tentative identification. Two species of echinoids are identified from rare disarticulated plates, namely Melonechinus? sp. (ambulacral plates) and archaeocidarid sp. (interambulacral plate); more and superior material will be necessary to confirm this division. Crinoids include a cladid brachial ossicle sp. with a distinctive sculpture; a columnal of a monobathrid camerate platycrinitid sp.; Annulocolumnus (col.) sp. cf. A. annulus Donovan, a columnal morphotaxon with an unusually broad axial canal; and Cyclocyclicus (col.) spp. This is the tenth echinoid site to be recognized from the Mississippian of the White Peak. Examination of etched float blocks provides a further method of investigation of the echinoderm fauna of this and other limestone areas.     

First crinoid crown from the Permian Khuff Formation (Wordian) of Oman

Crinoidal debris is common from the basal parts of the open marine Khuff Formation in Oman; yet, little is known about the diversity and affinities of this fauna. Exallocrinus khuffensis n. gen., n. sp. is described from this unit, and is the first crinoid crown from outcrops of the Wordian, Lower Khuff Member, in the northern Huqf region of Oman. This new crinoid is among the youngest Paleozoic crinoids known, yet it has a combination of more stemward and crownward characters. Because of the uncertainties concerning the latest Paleozoic and earliest Mesozoic crinoid phylogeny, Exallocrinus n. gen. is questionably assigned to the Ampelocrinidae.     

Crinoids columnals (Echinodermata) of the Ererê Formation (late Eifelian–early Givetian,Amazon Basin), State of Pará, Brazil

The faunal composition of stalked echinoderms in the Brazilian Devonian is still largely unknown despite the great abundance of crinoids in the shallow epicontinental seas of the Paleozoic. The first Devonian crinoids of Brazil, recorded in the literature in 1875 and 1903, were from the sedimentary rocks of the Ererê Formation in the Amazon Basin. Since then, the echinoderms of this formation have not been studied. This study, based on isolated pluricolumnals and columnals, described and identified Botryocrinus meloi n. sp., the first record for this genus in Brazil. In addition to this species, two other morphological patterns were identified: Tjeecrinus sp. and Morphotype AM/Er-01. The form of occurrence of the crinoid material and the paleoautoecology of B. meloi allow preliminary characterization of the habitat as a moderately deep water with weak to moderate currents and soft substrate. The similarity between B. meloi and Botryocrinus montguyonensis and of Tjeecrinus? sp. and T. crassijugatus, from the Devonian of the Armorican and Rhenan Massif, represents new evidence for the existence of contact between the faunas of the Amazon Basin with those of northern Gondwana and Armorica during the Middle Devonian.     

Neoichnology of Chalk cobbles from north Norfolk,England: implications for taphonomy and palaeoecology

The study of marine biota of the North Sea from the perspective of its relevance to palaeontology (Aktuopaläontologie) has enabled modern patterns and processes to be identified that can also be recognised in ancient communities. A collection of Chalk cobbles from the coast of north Norfolk, eastern England, preserves a limited ichnofauna: Gastrochaenolites ornatus Kelly and Bromley; Gastrochaenolites isp.; and Caulostrepsis isp. All cobbles show evidence of physical modification (corrasion) since they were bored; no Gastrochaenolites is complete and no clast has Caulostrepsis on all sides, therefore clasts have been corraded subsequent to boring. Only one producing organism in Gastrochaenolites, probably Barnea sp., is preserved in situ. Gastrochaenolites borings were infested by a range of encrusting organisms after the demise (and removal of shells) of the boring bivalves: thread-like algae; agglutinated worm tubes; cheilostome bryozoan Cryptosula pallasiana (Moll); and serpulids Hydroides norvegica Gunnerus and Pomatoceras triqueter (Linnaeus). Such a suite of encrusting organisms would be impossible to see in a fossil boring filled with lithified sedimentary rock unless it was released by dissolution of the substrate. Such cobbles, corraded since they were infested by borers, leave only portions of traces for identification, commonly only to ichnogenus.     

The terrestrial fauna of the Late Triassic Pant-y-ffynnon Quarry fissures,South Wales,UK and a new species of Clevosaurus (Lepidosauria: Rhynchocephalia)

Pant-y-ffynnon Quarry in South Wales yielded a rich cache of fossils in the early 1950s, including articulated specimens of new species (the small sauropodomorph dinosaur Pantydraco caducus and the crocodylomorph Terrestrisuchus gracilis), but no substantial study of the wider fauna of the Pant-y-ffynnon fissure systems has been published. Here, our overview of existing specimens, a few described but mostly undescribed, as well as freshly processed material, provides a comprehensive picture of the Pant-y-ffynnon palaeo-island of the Late Triassic. This was an island with a relatively impoverished fauna dominated by small clevosaurs (rhynchocephalians), including a new species, Clevosaurus cambrica, described here from a partially articulated specimen and isolated bones. The new species has a dental morphology that is intermediate between the Late Triassic Clevosaurus hudsoni, from Cromhall Quarry to the east, and the younger C. convallis from Pant Quarry to the west, suggesting adaptive radiation of clevosaurs in the palaeo-archipelago. The larger reptiles on the palaeo-island do not exceed 1.5?m in length, including a small carnivorous crocodylomorph, Terrestrisuchus, and a possible example of insular dwarfism in the basal dinosaur Pantydraco.     

Early Cretaceous (Valanginian) sea lilies (Echinodermata, Crinoidea) from Poland

Valanginian strata in central epicratonic Poland have recently yielded crinoids, not previously recorded from the area. The fauna comprises isocrinids (Balanocrinus subteres, B. gillieroni, “Isocrinus?” lissajouxi), millericrinids (Apiocrinites sp.) and comatulids (Comatulida indet.). For comparison, a few samples of isocrinids from Valanginian strata of Hungary (Tethyan province) were also analysed. The isocrinids, cyrtocrinids and roveacrinids (sensu Rasmussen 1978 inclusive of Saccocoma sp.) were already known from the Valanginian of the southernmost Tethyan regions of Poland (Pieniny Klippen Belt and Tatra Mountains). The current study demonstrates their occurrence in central epicratonic Poland, and suggests that many Jurassic to Cretaceous stalked crinoid taxa (mainly isocrinids) predominated in the shallow-water settings of this area. Thus, the hypothesis of migration (at least from mid-Cretaceous onwards) to deep-water areas, as a response to an increase of the number of predators during the Mesozoic marine revolution, seems not to be universally applicable.     

A primitive cladid crinoid from the Jiacun Group,Tibet (Darriwilian,Middle Ordovician)

A collection of numerous crinoid pluricolumnals from the uppermost Darriwilian (Middle Ordovician) of Tibet were derived from one biological species of crinoid. The specimens were collected from well‐lithified, bioclastic shelf limestones of the upper part of the Lower Chiatsun Group, Pygodus serra Biozone; coeval rocks of similar lithology outcrop at the summit of Everest. A new crinoid morphospecies, Segmentocolumnus (col.) hanshessi, is tentatively considered a ‘stem‐group cladid’, perhaps a dendrocrinid. The proxistele is broad and pentagonal in section with a broad, pentagonal axial canal; the mesistele of similar gross morphology is more slender with a regularly heteromorphic column and a similarly wide axial canal; the dististele is a terminal dendritic radice with a pentastellate axial canal. In the mesistele, the meric sutures correspond to the centres of the sides of the column, but in the dististele they occur in the angles. This range of morphologies would have led to their inclusion in at least two morphogenera if they had not been closely associated; as they belong to a single biological species, they have been ‘lumped’ together herein. This is a rare contribution to our knowledge of the early crinoids from a region outside Europe and North America. Copyright © 2012 John Wiley & Sons, Ltd.     

Bathonian (Middle Jurassic) crinoids of the Hidas Valley (Mecsek Mountains, S Hungary) and their biogeographic significance

The Bathonian crinoid fauna that occurs in red nodular limestone and argillaceous limestones from the Hidas Valley, Mecsek Mts (southern Hungary) consists of three isocrinid and six cyrtocrinid species. Isocrinids are represented by Balanocrinus inornatus (d’Orbigny), B. berchteni Hess and Pugin and Balanocrinus sp. Cyrtocrinids are represented by Phyllocrinus stellaris Zar?czny, P. birkenmajeri G?uchowski, P. malbosianus d’Orbigny, Apsidocrinus sp., Lonchocrinus sp., and the new species Psalidocrinus hidasinus sp. nov. This last species is the earliest occurrence of the genus Psalidocrinus previously known from the Early Tithonian to Valanginian. This is the first crinoid fauna described from the Middle Jurassic (Bathonian) of Hungary. The co-occurrence of isocrinids and cyrtocrinids indicates an environment subject to weak currents. The stratigraphical and geographical distribution of the identified cyrtocrinid genera and species suggests a Tethyan origin and subsequent migration to the northern Tethyan shelf.     

Late Tournaisian conodonts from the Taungnyo Group near Loi Kaw,Myanmar (Burma): Implications for Shan Plateau stratigraphy and evolution of the Gondwana-derived Sibumasu Terrane

Carboniferous conodonts are reported for the first time from Myanmar (Burma). Conodont faunas representative of the Scaliognathus anchoralis and Gnathodus typicus–Protognathodus cordiformis conodont zones date the sampled Taungnyo Group south of Loi Kaw, Kayah State as late Tournaisian confirming a Mississippian (Lower Carboniferous) age for the sampled part of this stratigraphic unit. The dated strata are stratigraphically just below the Tournaisian–Visean (T-V) boundary. Tournaisian strata are thus for the first time unequivocally demonstrated in the Shan Plateau region of Myanmar. Similar conodont faunas from the T-V boundary interval in SE Asia indicate a complete stratigraphic sequence at this level in shallow-marine sequences on intra-Tethyan Cathaysian tectonic blocks (South China) and in deep-marine Palaeo-Tethyan sediments (cherts of the Inthanon suture zone, Thailand). However, in shallow-marine sequences on the Sibumasu Block, located on the NE margin of Gondwana in the Carboniferous, they demonstrate a non-sequence or unconformity at this level, also seen elsewhere in Gondwana. Biogeographic links between upper Tournaisian and early Visean conodonts on the Sibumasu Terrane and Laurentia and Eastern Australian Gondwana support a NW Australian Gondwana margin position for Sibumasu in the Late Palaeozoic.     

Utility of crinoid columnals in palaeontology illustrated by a new species: Clare Shale Formation (Carboniferous), Doolin,County Clare,western Ireland

Well-preserved specimens, such as complete individuals, crowns and cups, are the common focus for crinoid systematic research. Yet the majority of specimens are disarticulated ossicles which are essentially ignored. The incompleteness of the fossil record is even more so when we ignore potential sources of data. A new species of crinoid comes from a monospecific assemblage from the Pennsylvanian (Upper Carboniferous) of western Ireland. All specimens are from a float block of the Clare Shale Formation (Bashkirian stage) at Fisherstreet Bay, Doolin, County Clare, western Ireland. Heloambocolumnus (col.) harperi gen. et sp. nov. has a pentagonocyclic, heteromorphic column; the small, central lumen is in a shallow, circular claustrum; the articulation is radial symplectial; the crenulae are slightly swollen and peg-like close to the circumference; nodals have rounded, unsculptured epifacets; nodal articular facets are sunken and in which narrow internodals are situated; and circlets of tubercles on epifacet surround priminternodals. These columnals are associated with robust, uniserial brachial ossicles. This crinoid is most likely a cladid.     

Echinoids as hard substrates: varied examples from the Oligocene of Antigua,Lesser Antilles

A collection of unremarkably preserved fossil irregular echinoids from the Upper Oligocene (Chattian) Antigua Formation of Antigua, Lesser Antilles, nonetheless provides evidence of a range of palaeoecological interactions. A dead test of the heart urchin Eupatagus sp. formed a hard substrate for the attachment of gregarious Thecidellina? sp., a thecidoid brachiopod. Although obligate encrusters, these brachiopods more commonly occur as disarticulated valves free of the substrate in the Antillean fossil record. Elongate pits in test fragments were formed, variously, before and after the death of the host echinoids. These depressions on the external surface were formed either by invertebrates excavating domiciles or by claws or teeth; the echinoid later reclaimed the pits and grew new tubercles in the base. Post-mortem pits lack such new tuberculation. A test of Eupatagus sp. bears the boring Oichnus isp., formed either by a predator (gastropod?) or after the death of the echinoid (domicile), and a serpulid worm tube which grew on the test subsequent to the echinoid’s death. The echinoid fauna of the Antigua Formation has been easy to collect and specimens are to be found in many museums; they now await re-examination to reveal palaeosynecological data analogous to that determined from the fragments discussed herein.     

Taphonomy and palaeoecology of the gastropod fauna from a Late Cretaceous rocky shore, Sweden

A gastropod fauna comprising 17 species, each represented by a limited number of specimens, is described from a Late Cretaceous, late early Campanian rocky shore at Ivö Klack, southern Sweden. The gastropod fauna is associated with the most diverse ancient rocky shore fauna ever found. However, the low gastropod species diversity compared to the faunas of modern rocky shores is ascribed to taphonomic factors, notably dissolution of the aragonitic shells, but the predominance of epifaunal herbivores is indicative of a guild structure similar to that found on modern rocky shores. The presence of drill holes assigned to the ichnospecies Oichnus simplex suggests the former presence of muricid gastropods which have not been found as body fossils. A single drill hole is referred to Oichnus paraboloides and was probably made by a naticid gastropod. The infaunal mode of life of naticids makes preservation of such drill holes difficult, since the majority of infaunal prey such as burrowing bivalves has aragonitic shells which are not preserved. The relatively high number of species in comparison to many other Late Cretaceous rocky shore faunas, offers an opportunity to compare gastropod guild structure at Ivö Klack with modern counterparts, even though taphonomic processes such as mechanical destruction and aragonite dissolution have played an important role in the fossil gastropod assemblage.     

吉林省中部早石炭世杜内期的腕足动物化石

描述的化石是采自吉林省中部杜内期北通气沟组,共17个属24个种,包括3个新种。这个动物群的特点与我国南方杜内期动物群的特点不同,而与美国密西西比系金德胡克组、苏联库茨纳兹盆地和莫斯科盆地杜内期的动物群非常相像。类似的动物群在新疆、甘肃和内蒙等省和自治区均已发现。吉中地区杜内期腕足动物群,属于北美—西伯利亚古生物地理区系。     

Ammonites (Phylloceratina,Lytoceratina and Ancyloceratina) and organic-walled dinoflagellate cysts from the Late Barremian in Boljetin,eastern Serbia

Late Barremian ammonite fauna from the epipelagic marlstone and marly limestone interbeds of Boljetin Hill (Boljetinsko Brdo) of Danubic Unit (eastern Serbia) is described. The ammonite fauna includes representatives of three suborders (Phylloceratina, Lytoceratina and Ancyloceratina), specifically Hypophylloceras danubiense n. sp., Lepeniceras lepense Rabrenović, Holcophylloceras avrami n. sp., Phyllopachyceras baborense (Coquand), Phyllopachyceras petkovici n. sp., Phyllopachyceras eichwaldi eichwaldi (Karakash), Phyllopachyceras ectocostatum Drushchits, Protetragonites crebrisulcatus (Uhlig), Macroscaphites perforatus Avram, Acantholytoceras cf. subcirculare (Avram), Dissimilites cf. trinodosus (d'Orbigny) and Argvethites? sp. The taxonomic composition and percent abundance of the identified ammonites indicate that their taxa are predominantly confined to the Tethyan realm. Ammonites with smooth and slightly sculptured shells predominate among the studied fauna. The ammonite-bearing succession from Boljetin represents the lower part of the Upper Barremian, ranging in ammonite zonation from the Toxancyloceras vandenheckei Zone to the lower part of the Imerites giraudi Zone. The associated organic-walled dinoflagellate cysts confirm the Late Barremian age of the ammonite-bearing levels.