Revision of Strunian crinoids and blastoids from Germany

In 1930W.E. Schmidt described all known crinoids from the German Early Carboniferous, including the Etroeungt beds of Germany, which are now judged to be latest Devonian (Famennian) in age. On a global basis, Famennian camerate crinoids generally show a closer relationship to succeeding Early Carboniferous faunas than they do to older Frasnian or Middle Devonian crinoids, which also is the case for the Strunian fauna. Holdovers from older Devonian faunas include, among others,Adelocrinus, a descendant of olderArthroacantha, in England and Germany, andPetaloblastus, which is one of the youngest genera of the blastoid family Hyperoblastidae. Precursors of younger Early Carboniferous groups include platycrinoids, primitive actinocrinoids, dichocrinoids, and the blastoid genusDoryblastus, which is one of the oldest members of the family Orbitremitidae. All of these groups, which became important parts of the Early Carboniferous crinoid and blastoid radiation, give Famennian crinoid faunas much more of an Early Carboniferous than a Devonian aspect. Rhipidocrinus schmidti n. sp. is erected for specimens that originally were reported from the Etroeungt asRhodocrinus uniarticulatus. We judge that there are currently four valid species assigned toRhipidocrinus: R. crenatus, R. perloricatus, R. praecursor, and our new species,R. schmidti. Hydriocrinus ratingensis Schmidt is reassigned to ?Sostronocrinus. We note thatSchmidt (1906), notJaekel (1906) as has been reported previously, is the author ofRhipidocrinus perloricatus. Owing to the poor preservation of the Etroeungt material, we regard the namePlatycrinites wunstorfi Schmidt 1930 to be a nomen nudum.     

Late Famennian gastropoda from south-west England

The gastropod fauna of the Upper Devonian Baggy and Pilton formations in south‐west England is revised and includes some 30 taxa. The topmost part of the Upper Famennian succession in Devon is represented by clastic near‐shore and shallow shelf sediments, indicating a short‐term transgressive phase (‘Strunian Transgression’). The sequence yields a highly diverse fauna dominated by brachiopods and ostracodes, locally supplemented by crinoids, bryozoans, trilobites and molluscs. The taxa ‘Patellostium’britannicum sp. nov., Angyomphalus (Angyomphalus) junius sp. nov. and Dictyotomaria eurocapillaria sp. nov. are erected; a junior homonym is replaced by Macrochilina? piltonensis nom. nov. The gastropod fauna displays an independent character, where latest Devonian faunal elements overlap with Late Palaeozoic taxa expressing a transition similar to that of the bivalves, brachiopods, echinoderms and corals, without a sharp faunal break at the Devonian/Carboniferous boundary. Apart from the Caenogastropoda, all subclasses of gastropods are represented. Members of the bellerophontoids, pleurotomarioids and loxonematoids are most abundant, followed by murchisonioids, naticimorphs, euomphalomorphs and platyceratoids. The various gastropod groups represent different ecological demands and trophic categories, and together with the accompanying fauna indicate that nearly all habitats and niches were occupied in the shallow South Laurussian Shelf.     

Tube foot preservation in the Devonian crinoid Codiacrinus from the Lower Devonian Hunsrück Slate,Germany

Fossilized tube feet are described on Codiacrinus schultzei Follmann from the Lower Devonian Hunsrück Slate of Germany. This is the first definitive proof of tube feet on any fossil crinoid. Three lightly pyritized, flattened tube feet are preserved in a single interray of this cladid crinoid. The tube feet were at least 7 mm long. Their preservation is very similar to the tube feet reported previously from a Hunsrück ophiuroid, except that the Codiacrinus tube feet have small papillae, similar to living crinoids.     

New Pennsylvanian (Moscovian) crinoids from the Qijiagou Formation,Taoshigo Valley,Xinjiang Uyghur Autonomous Region,Western China

New collections and revision of previously collected Moscovian crinoids from the Qijiagou Formation of the Taoshigo Valley near Turpan, Xinjiang Uyghur Autonomous Region, Western China, add to the generic diversity of the fauna. This camerate-rich echinoderm fauna is now recognized as containing at least one blastoid, five camerate, and ten cladid crinoid genera. The fauna shows greatest affinity at the family level with Moscovian crinoid faunas of Japan and North America.New taxa proposed are: Rhepocatillocrinus tianshanensis n. gen. n. sp.; Binariacrinus alveus n. gen. n. sp.; Bassocrinus abyssus n. gen. n. sp.; and Brabeocrinus asiaensis n. sp.     

Aspects of crinoid palaeontology of the North Esk Inlier,Scotland (Silurian,Llandovery, Telychian)

Abstract: The described fauna of well‐preserved Llandovery (Telychian) echinoderms from the North Esk Inlier, including six crinoids, one echinoid and seven starfish species, is mainly allochthonous. Most of these taxa are known only from starfish beds, channel fill deposits probably representing submarine mass flows and preserving a biota probably derived from elsewhere, presumably shallower water. Only one crinoid species, Pisocrinus cf. campana Miller, is recognized as a common fossil away from the starfish beds and is a biostratigraphic marker for the base of the Wether Law Linn Formation, forming part of the Skenidioides‐Cyrtia Association. Crinoid columnals preserved perpendicular to bedding (that is, in putative life position) in Lamont’s bivalve bed, Deerhope Formation, are tentatively interpreted as being in situ by comparison with a similar occurrence in the Silurian of Arisaig, Nova Scotia. Two new species of crinoid are described, the cladid Dendrocrinus? sp. and the columnal morphospecies Pentagonocyclicus (col.) lamonti sp. nov.     

Stenolaemate bryozoans from the Geirud Formation (Upper Devonian/Lower Carboniferous) of Central Alborz (Iran)

Two bryozoan species are described from the Geirud Formation (Upper Devonian/Lower Carboniferous) of Central Alborz (Iran). Trepostome Schulgina mutabilis Troizkaya, 1975 is known from the Upper Devonian (Famennian) of Central Kazakhstan. The new species Ascopora geirudensis n. sp. is the earliest known representative of the rhabdomesine genus Ascopora Trautschold, 1876.     

Anatomical reinvestigation of Archaeopteris macilenta from the Upper Devonian (Frasnian) of South China

Abstract Archaeopteris macilenta is one of the most widespread plants in the Late Devonian. Based on fossils from the Frasnian Huangjiadeng Formation, Yichang District of Hubei Province, for the first time we study in detail the anatomy of this progymnosperm plant in South China. Ultimate axes are protostelic with three xylem sympodia and lack secondary tissue. Penultimate axes are eustelic, bearing eight sympodia and a thin band of secondary xylem. Radially symmetrical sympodia of mesarch primary xylem produce traces of appendages in a spiral arrangement. Archaeopteris macilenta and A. halliana (A. roemeriana) are dominant in the Frasnian and Famennian, respectively. Comparisons with these two species from other tectonic plates indicate consistent stelar architectures. Global spread, continuous occurrence, and identical anatomy during the Late Devonian indicate that Archaeopteris survived the Frasnian–Famennian extinction event. In this time, endemic genera and cosmopolitan taxa, including Archaeopteris, suggest the palaeogeographic isolation of South China and certain associations with other plates.     

NEW CLADID AND FLEXIBLE CRINOIDS FROM THE MISSISSIPPIAN (TOURNAISIAN, IVORIAN) OF ENGLAND AND WALES

Abstract:  The modern study of fossil crinoids began with J. S. Miller who, in 1821, described specimens from southern England, nearby Wales and other regions, and named several common Early Carboniferous genera. Later, in 1950–60, James Wright monographed all known Early Carboniferous crinoids from the British Isles. In spite of such previous scrutiny, we recognize here two new genera among species already described: Glamorganocrinus gen. nov. (type species: Ophiurocrinus gowerensis Wright, 1960) from South Wales and Mendipocrinus gen. nov. (type species: Poteriocrinus latifrons Austin and Austin, 1847) from southern England. These new genera increase the number of advanced cladid genera in the Ivorian Substage of the Tournaisian in western Europe to 18, and the total number of crinoid genera to 36. A review of species assigned to Mespilocrinus has led to the recognition of M. granulifer De Koninck and LeHon, 1854 as a nomen dubium. A new species of Mespilocrinus , M. wrighti sp. nov., is described from the Ivorian of South Wales; this is the most highly derived species of the genus, as based on a phylogenetic analysis including ten species and 13 characters, with Pycnosaccus as the outgroup. A single, well-ordered tree resulted from this analysis. Interpretation of this tree suggests that the centre of evolution for Mespilocrinus was North America, where three species appeared during the Kinderhookian (early Tournaisian), rapidly achieving morphological disparity within the genus. This radiation event was part of the overall explosive radiation of crinoids following the Late Devonian mass extinction event when crinoid diversity was at a global minimum during the Frasnian. Recovery began during the Famennian, followed by an explosive radiation in the Tournaisian.     

Biodiversity and historical biogeography of stalked crinoids (Echinodermata) in the deep sea

About 95 species of stalked crinoids are now described from 60m to hadal depths, but our knowledge remains far from complete. Depending on which species concept is used, estimates of species richness can be dramatically different. It is necessary to have a homogeneous concept for taxonomic units. The abundance of the crinoid fossil record allows a discussion of the ancestry of deep sea crinoid fauna. Stalked crinoids have a horizontal diversity pattern with three regional centres of high diversity (i.e. western tropical Pacific, western tropical Atlantic and north-eastern Atlantic). Vertical patterns show two faunal strata which vary in importance among provinces. The epibathyal stratum has apparently remained relatively similar in intertropical areas since the Mesozoic. Despite environmental changes related to glaciation since the Middle Miocene, the deepest crinoid fauna (i.e. the deep sea fauna sensu stricto at depths more than 1000 ± 200 m) have a very ancient origin with a dispersion closely related to plate tectonics. The bathyal fauna on hard substrates includes a few living fossils and has a high historical interest.     

New discoveries of tetrapods (ichthyostegid‐like and whatcheeriid‐like) in the Famennian (Late Devonian) localities of Strud and Becco (Belgium)

The origin of tetrapods is one of the key events in vertebrate history. The oldest tetrapod body fossils are Late Devonian (Frasnian–Famennian) in age, most of them consisting of rare isolated bone elements. Here we describe tetrapod remains from two Famennian localities from Belgium: Strud, in the Province of Namur, and Becco, in the Province of Liège. The newly collected material consists of an isolated complete postorbital, fragments of two maxillae, and one putative partial cleithrum, all from Strud, and an almost complete maxilla from Becco. The two incomplete maxillae and cleithrum from Strud, together with the lower jaw previously recorded from this site, closely resemble the genus Ichthyostega, initially described from East Greenland. The postorbital from Strud and the maxilla from Becco do not resemble the genus Ichthyostega. They show several derived anatomical characters allowing their tentative assignment to a whatcheeriid‐grade group. The new tetrapod records show that there are at least two tetrapod taxa in Belgium and almost certainly two different tetrapod taxa at Strud. This locality joins the group of Devonian tetrapod‐bearing localities yielding more than one tetrapod taxon, confirming that environments favourable to early tetrapod life were often colonized by several tetrapod taxa.     

Unusual tabulate‐crinoid biocoenosis from the Lower Devonian of Morocco

The Early Devonian (Pragian: sulcatus to pireneae conodont zones) crinoid–coral biocoenosis from Hamar Laghdad, Morocco contains fragments of crinoid stalks of various taxa encrusted by spherical and ellipsoidal coralla of the tabulate coral Hamarilopora minima. These corals were encrusting host crinoids syn vivo, and this is evidenced by pluricolumnals exceeding 30 elements overgrown from all sides. Most known to date crinoid–epibiont associations display various types of reaction to the epibiont, such as swellings and deformations. In the case discussed here, no clear interaction is visible; therefore, this association can be classified as paroecia. It can be inferred, however, that due to a change in mechanical properties of the crinoid stalk (losing flexibility), the epizoan influence on the host was negative, while the coral was profiting from the elevated position over the seafloor and nutrient‐bearing water currents. It can be supposed that this interaction was close to parasitism. No strict species‐specific relationship between the epizoan and the host was observed.     

New Late Devonian strophomenids (Brachiopoda) from Mongolia

Two new strophomenid species, Leptagonia barunkhuraica sp. nov. (Famennian) and Floweria mongolica sp. nov. (Frasnian), from the Upper Devonian of the Barunkhurai Depression of southern Mongolia are described.     

Infestation of Middle Devonian (Givetian) camerate crinoids by platyceratid gastropods and its implications for the nature of their biotic interaction

One of the classic examples of biotic interactions preserved in the fossil record is that between crinoids and infesting platyceratid gastropods. This relationship, spanning an interval from the Middle Ordovician to the end of the Permian, is recognized by the firm attachment and positioning of platyceratids over the anal vent of their hosts. Several hypotheses have been proposed to explain this interaction; the most widely accepted is that the gastropods were coprophagous commensals, feeding on crinoid excrement without any significant detriment to their hosts. The purpose of this investigation was to test this hypothesis. Two species of Middle Devonian camerate (Monobathrida, Compsocrinina) crinoids, Gennaeocrinus variabilis Kesling & Smith 1962 and Corocrinus calypso (Hall 1862), were used in this investigation. The data consisted of 426 individuals of G. variabilis collected near Rockport, Michigan, 30 of which were infested, and 188 individuals of C. calypso collected near Arkona, Ontario, Canada, of which 25 were infested. Length and volume were measured for each crinoid to determine whether a significant difference existed in the size of infested versus uninfested individuals. The results indicated that for both species of crinoids individuals infested by snails were significantly smaller than uninfested individuals (p < 0.05). We explored a variety of scenarios to explain this pattern and conclude that they falsify the null hypothesis that the crinoid-gastropod relationship was strictly commensal. The smaller size of the infested crinoids is interpreted as a consequence of nutrient-stealing by the parasitic gastropods, a strategy that finds analogs in modern seas. Moreover, the absence of platyceratids on the largest crinoids suggests that large size may have inferred immunity from lasting infestation.     

The fate of the homoctenids (Tentaculitoidea) during the Frasnian–Famennian mass extinction (Late Devonian)

The homoctenids (Tentaculitoidea) are small, conical‐shelled marine animals that are among the most abundant and widespread of all Late Devonian fossils. They were a principal casualty of the Frasnian–Famennian (F‐F, Late Devonian) mass extinction, and thus provide an insight into the extinction dynamics. Despite their abundance during the Late Devonian, they have been largely neglected by extinction studies. A number of Frasnian–Famennian boundary sections have been studied, in Poland, Germany, France, and the USA. These sections have yielded homoctenids, which allow precise recognition of the timing of the mass extinction. It is clear that the homoctenids almost disappear from the fossil record during the latest Frasnian ‘Upper Kellwasser Event’. The coincident extinction of this pelagic group, and the widespread development of intense marine anoxia within the water column, provides a causal link between anoxia and the F‐F extinction. Most notable is the sudden demise of a group, which had been present in rock‐forming densities, during this anoxic event. One new species, belonging to Homoctenus is described, but is not formally named here.     

A Lower Devonian hexacrinitid crinoid (Camerata,Monobathrida) from south-west England

The most species-rich and widespread crinoid clade in the type area of the Devonian of south-west England is the monobathrid camerate family Hexacrinitidae Wachsmuth and Springer. These crinoids occur either as thecae (Middle Devonian) or pluricolumnals and columnals (Lower to Upper Devonian). The first new, nominal species of hexacrinitid, probably Oehlerticrinus Le Menn, to be described from this region since the nineteenth century is Oehlerticrinus peachi sp. nov. from the Lower Devonian Looe Basin of southern Cornwall. This specimen is mouldic and somewhat flattened, retaining the proxistele and arms. Diagnostic features include the heteromorphic proxistele with circlets of long, unbranched radices directed towards the crown; the high, box-like (=skyphosiform) theca with a flattened base; a thecal plate sculpture of tubercles and ridges arranged in triangles; and the pinnulate arms. Any uncertainty in identifying this species at the generic level rests with the arms being pinnulate, not ramulose as is common in Hexacrinites Austin and Austin, indicating that O. peachi is closer to Oehlerticrinus Le Menn.     

New bryozoans from the Devonian of Gorny Altai (Russia)

New bryozoans Saffordotaxis altaicus sp. nov. from the Givetian Stage (Middle Devonian) and Cyphotrypa olgae sp. nov. and Crustopora aliena sp. nov. from the Famennian Stage (Upper Devonian) are described from the Devonian of Gorny Altai.     

A fish and tetrapod fauna from Romer's Gap preserved in Scottish Tournaisian floodplain deposits

The end‐Devonian mass extinction has been framed as a turning point in vertebrate evolution, enabling the radiation of tetrapods, chondrichthyans and actinopterygians. Until very recently ‘Romer's Gap’ rendered the Early Carboniferous a black box standing between the Devonian and the later Carboniferous, but now new Tournaisian localities are filling this interval. Recent work has recovered unexpected tetrapod and lungfish diversity. However, the composition of Tournaisian faunas remains poorly understood. Here we report on a Tournaisian vertebrate fauna from a well‐characterized, narrow stratigraphic interval from the Ballagan Formation exposed at Burnmouth, Scotland. Microfossils suggest brackish conditions and the sedimentology indicates a low‐energy debris flow on a vegetated floodplain. A range of vertebrate bone sizes are preserved. Rhizodonts are represented by the most material, which can be assigned to two taxa. Lungfish are represented by several species, almost all of which are currently endemic to the Ballagan Formation. There are two named tetrapods, Aytonerpeton and Diploradus, with at least two others also represented. Gyracanths, holocephalans, and actinopterygian fishes are represented by rarer fossils. This material compares well with vertebrate fossils from other Ballagan deposits. Faunal similarity analysis using an updated dataset of Devonian–Carboniferous (Givetian–Serpukhovian) sites corroborates a persistent Devonian/Carboniferous split. Separation of the data into marine and non‐marine partitions indicates more Devonian–Carboniferous faunal continuity in non‐marine settings compared to marine settings. These results agree with the latest fossil discoveries and suggest that the Devonian–Carboniferous transition proceeded differently in different environments and among different taxonomic groups.     

The oldest Gondwanan cephalopod mandibles (Hangenberg Black Shale,Late Devonian) and the mid‐Palaeozoic rise of jaws

It is widely accepted that the effects of global sea‐level changes at the transition from the Devonian to the Carboniferous are recorded in deposits on the shelf of northern Gondwana. These latest Devonian strata had been thought to be poor in fossils due to the Hangenberg mass extinction. In the Ma'der (eastern Anti‐Atlas), however, the Hangenberg Black Shale claystones (latest Famennian) are rich in exceptionally preserved fossils displaying the remains of non‐mineralized structures. The diversity in animal species of these strata is, however, low. Remarkably, the organic‐rich claystones have yielded abundant remains of Ammonoidea preserved with their jaws, both in situ and isolated. This is important because previously, the jaws of only one of the main Devonian ammonoid clades had been found (Frasnian Gephuroceratina). Here, we describe four types of jaws of which two could be assigned confidently to the Order Clymeniida and to the Suborder Tornoceratina. These findings imply that chitinous normal‐type jaws were likely to have already been present at the origin of the whole clade Ammonoidea, i.e. in the early Emsian (or earlier). Vertebrate jaws evolved prior to the Early Devonian origin of ammonoids. The temporal succession of evolutionary events suggests that it could have been the indirect positive selection pressure towards strong (and thus preservable) jaws since defensive structures of potential prey animals would otherwise have made them inaccessible to jawless predators in the course of the mid‐Palaeozoic marine revolution. In this respect, our findings reflect the macroecological changes that occurred in the Devonian. [Correction added on 28 July 2016 after first online publication: In the Abstract, the sentence “Vertebrate jaws probably … in the Early Devonian” was amended]     

Devonian (Eifelian) disparid crinoids from the Zeravashan range,southern Tien Shan,Uzbekistan

No Devonian disparid crinoids have been described from Uzbekistan, although parahexacrinid thecae and assorted columnals have been described in several papers in the past 50 years. Discovery of a cup of Pisocrinus and a thecae of Haplocrinites are the first of these genera known from Uzbekistan. The stratigraphic range of Haplocrinites is revised to be late Early Devonian (Emsian) to Early Carboniferous (Tournaisian) because Silurian species assigned to the genus are based on loose ossicles that morphologically do not belong to the genus. The paleogeographic ranges of Pisocrinus and Haplocrinites are extended into Uzbekistan. Haplocrinites uzbekistanensis n. sp. is described.     

Assessment of hidden diversity of crinoids and their symbionts in the Bay of Nhatrang,Vietnam

Crinoid associates represent an abundant and diverse, but poorly explored, component of the hidden biodiversity of coral-reef ecosystems. We studied data from 5 years of collecting in the Bay of Nhatrang (BN), Vietnam, to assess the diversity of crinoids and their symbionts, to compare it with other areas of the Indo-West Pacific, and to elucidate the extent to which the observed diversity of crinoids and their symbionts corresponds to their true diversity. In total, about 2,287 specimens of symbionts belonging to 70 species were found on 203 specimens of crinoids belonging to 33 species. Among the crinoids, the most numerous species were Himerometra robustipinna (36 specimens) and Cenometra bella (29 specimens), among the symbionts the polychaete Paradyte crinoidicola (c. 850 specimens) and the galatheid crustacean Allogalathea elegans (180 specimens). Species accumulation curves suggest that we have sampled most of the crinoid diversity in the BN, whereas the diversity of their symbionts remained undersampled. Estimated species richness of crinoids was higher than previously observed richness, and varied from 39 (estimated by bootstrap) to 46 (jackknife 2). Estimated species richness of symbionts was higher than observed richness, and varied from 71 (bootstrap) to 93 (jackknife 2). We suggest a slight increase in the number of crinoid species to result from more detailed studies of nocturnal species, and an increase in the number of symbiotic species when studies of nocturnal crinoid associates and sibling species among decapods are included. Our study revealed a rather rich crinoid fauna in the bay compared to other areas of the Indo-West Pacific, and the highest species richness of crinoid associates known from anywhere in the World Ocean.