Clypeasteroid echinoid tests as benthic islands for gastrochaenid bivalve colonization: evidence from the Middle Miocene of Tarragona,north‐east Spain

Abstract: Middle Miocene tests of Clypeaster from L’Arrabassada (Tarragona, north‐east Spain) show evidence of intense endoskeletozoan colonization, preserved as borings and associated carbonate secretions that allow gastrochaenid bivalves to be identified as the colonizers. Two modes of occurrence have been recognized for these bivalve dwelling cavities; ‘intrastereom clavate borings’ which are restricted to the echinoid stereom, and ‘semi‐endoskeletal dwellings’, which penetrate across the test wall and extend as carbonate crypts into the sediment fill of the internal test cavity. Their size, density and position rule out a syn‐vivo relationship with the echinoids and demonstrate that colonization was post mortem. Because of the endurance of clypeasteroid skeleton and the pronounced bell‐shaped morphology of Clypeaster, the tests of these echinoids provided the most suitable substrates for hard‐bottom colonizers on an otherwise sandy seafloor. The scenario described from Tarragona can be extended to other Neogene and Quaternary strata elsewhere; there is ample evidence for the long‐term utilization of tests of Clypeaster by gastrochaenid bivalves in shoreface palaeoenvironments.     

Perisphinctes orbignyi n. sp. and accompanying perisphinctid assemblage from the Lower Oxfordian (Cordatum Zone,Cordatum Subzone) in southern Poland (Zalas,Krakow region)

The paper presents preliminary characteristics of a new perisphinctid assemblage from the Lower Oxfordian section of Zalas near Krakow, regarded as the classic one for southern Poland. The assemblage is derived from strata of the Cordatum Subzone, Cordatum Zone. Two major groups of perisphinctids are differentiated. The assemblage appears similar to those described by Neumann from Cetechowitz 〚12〛, Bourseau from ‘Terres noires’ of de Beauvoisin in France 〚2〛, and Gygi from Switzerland 〚9〛. It fills very well a gap in our knowledge of this fauna from upper parts of the Lower Oxfordian and further studies should make it possible to propose and define a new biostratigraphic unit. A new ammonite species Perisphinctes (Arisphinctes?) orbignyi n. sp. is described. Its name is given to commemorate the bicentenary of the birth of Alcide d’Orbigny and his contribution to the developments of stratigraphy of the Mesozoic, especially Jurassic strata.     

The oldest record of a boring foraminifer: Early Permian of New Mexico,USA

The bioeroding foraminifer Troglotella incrustans Wernli and Fookes (Bolletino della Societa Paleontologica Italiana 31, 1992, 95), is widely reported from Bajocian?, and Oxfordian to Lower Cenomanian (with a Late Jurassic acme) shallow‐water limestones of the Tethyan realm. A single specimen of a boring foraminifer, assigned to T. incrustans, has now been observed from the Lower Permian (Sakmarian) Community Pit Formation of the Doña Ana Mountains, New Mexico, USA. Surviving the end‐Permian mass extinction, T. incrustans might be a Lazarus taxon that persisted in refuges. This finding represents the oldest record of a foraminifer exhibiting an euendolithic way of life. Boring foraminifera have not been previously recorded from strata older than the Jurassic. Boring traces of potentially foraminiferan origin, however, have been already reported from the Lower Carboniferous (?Ordovician).     

Selective feeding by the echinoid,Evechinus chloroticus,and the removal of plants from subtidal algal stands in Northern New Zealand

Summary Feeding choice of the echinoid Evechinus chloroticus was examined for six fucoid and one laminarian species of algae. Three experiments were conducted to determine the algal choice by echinoids under controlled conditions. In the first experiment, the seven algal species were presented to echinoids in laboratory conditions. The second experiment had replicates of the algal species placed randomly on a subtidal rocky reef where echinoids were abundant and randomly dispersed. For the third experiment, which was also field-based, replicates of one highly-ranked species, Ecklonia radiata, were presented to naturally dispersed Evechinus. In addition, a series of controlled observations was used to examine the order in which echinoids removed algae from mixed species stands on subtidal boulders and to determine if this was related to the experimentally demonstrated choices of algal species.The results of the first two experiments showed that there were differences between algal species in the amount of material grazed by echinoids. Rankings of algal species from the field experiment were not correlated with rankings from the laboratory experiment. The order of removal of algal species from natural stands was correlated with the laboratory-based experimental rankings of algal species, but not with the rankings from the field-based experiment or with algal species availability. There were differences between algal species in their vulnerability to grazing by echinoids, as measured by regression analyses on the amount of material grazed from algal replicates vs. the number of attached echinoids. Within each species, echinoid numbers exerted a non-linear effect on the removal of algal material. In the third experiment, where only one species of algae was presented, the echinoids still distributed themselves non-randomly amongst replicates, aggregating on some samples.Data on the finer scale distribution of algal species over the entire subtidal reef on which these experiments and observations were conducted indicate that Evechinus are not often presented with a choice of adult plants of several different species in natural stands.The evidence from this study supports the conclusion that feeding preferences by echinoids are labile and do not clearly exert the major influence on the removal of plants from natural stands. Preference, as determined from experimental rankings of algal species, is only one of a number of factors which may affect the removal of algae by echinoids. Other important factors are the density of echinoids present, algal susceptibility to removal, and the distribution and abundances of the various algal species and echinoids relative to each other. It is suggested that algal life history characteristics may be unaffected by echinoids and that coevolutionary arguments are not appropriate for describing echinoid-algal interactions.     

Loriolella,a key taxon for understanding the early evolution of irregular echinoids

Loriolella Fucini is an enigmatic and poorly understood Early Jurassic sea-urchin that has been interpreted both as a regular and an irregular echinoid. The discovery of new and well-preserved material of the type species L. ludovicii (Meneghini) from the Upper Pliensbachian (Lower Jurassic) of Lombardy, Italy, has clarified the morphology of this genus for the first time. Loriolella resembles regular echinoids in having a large (almost certainly monocyclic) apical disc enclosing the periproct, and large primary interambulacral tubercles and spines. However, its peristome is extremely small with only vestigial buccal notches and auricles, and the ambulacra expand adorally forming cassiduloid-like phyllodes. Cladistic analysis suggests that Loriolella is the earliest known microstomatid irregular echinoid. Its unique mixture of primitive and derived characters is important for interpreting the initial steps by which irregular echinoids arose from diadematoid-like regular ancestors.     

A new Late Jurassic species of the rare synechodontiform shark, <Emphasis Type="Italic">Welcommia</Emphasis> (Chondrichthyes,Neoselachii)

A new Jurassic species of the very rare and incompletely known synechodontiform shark, Welcommia, is described. The new species, Welcommia cappettai, is represented only by a single tooth, precluding reconstruction of its dentition in detail. Nevertheless, this specimen provides sufficient information and characteristics to establish its taxonomic status. Welcommia cappettai n. sp. occurs in the middle Oxfordian (Upper Jurassic) of south-western Germany. This is the first unambiguous record and named species of Welcommia from the Late Jurassic, substantially reducing the rather large gap in the fossil record of this synechodontiform taxon. So far, two Welcommia species from the Lower Jurassic of Belgium and the Lower Cretaceous of southern France have been described. An additional, still unnamed species seemingly occurs in the Oxfordian of southern France. The new species has plesiomorphic and apomorphic characteristics and, probably, an intermediate dental pattern that tentatively enables reconstruction of evolutionary trends in the dentition of this shark from small and compact teeth with broad, almost triangular cusps, to mesio-distally lengthened teeth with elongated mesial heels resulting in an extremely extended mesial cutting edge in addition to more delicate cusp and cusplets in advanced forms. These differences might be related to improved feeding mechanisms. It is hypothesized that Welcommia was predominantly a component of the Mediterranean faunal province. The disappearance of Welcommia in the Early Cretaceous remains ambiguous and might be related to competition by other sharks, for example hexanchiforms, or might represent a collecting bias and/or taxonomic misidentification of isolated teeth.     

Indo-Pacific echinoids in the tropical eastern Pacific

The existing literature reports that only one species of Indo-Pacific echinoid (Echinometra oblonga), occurs in the eastern Pacific. In this study we confirm the presence of this species at Islas Revillagigedo and also report the presence of two species ofEchinothrix (a genus hitherto unknown outside the Indo-Pacific) at Isla del Coco and at Clipperton Island. We also present evidence from isozymes and from mitochondrial DNA sequences indicating that at least one individual ofDiadema at Clipperton may belong to a maternal lineage characteristic of the west Pacific speciesD. savignyi. These data are consistent with the hypothesis that the observed populations of Indo-Pacific echinoid species are recent arrivals to the eastern Pacific, as opposed to the view that they are relicts of Tethyan pan-tropical distributions.Echinothrix diadema, in particular, may have arrived at Isla del Coco during the 1982-1983 El Nifio. In addition to Indo-Pacific species, Clipperton, Isla del Coco and the Revillagigedos contain a complement of eastern Pacific echinoids. The echinoid faunas of these islands should, therefore, be regarded as mixtures of two biogeographic provinces. Though none of the Indo-Pacific species are known to have reached the coast of the American mainland, their presence at the offshore islands of the eastern Pacific suggests that, for some echinoids, the East Pacific Barrier is not as formidable an obstacle to migration as was previously thought.     

Hrabalicrinus zitti gen. et sp. nov., and other Upper Jurassic crinoids (Echinodermata,Crinoidea) from the Brno area (Czech Republic)

Jurassic (Oxfordian) crinoids from the Brno area (Czech Republic) are described in details for the first time. A rich crinoid assemblage consisting of cups, isolated cup elements, brachial plates, columnals, pluricolumnals, and cirrals is assigned to isocrinids Isocrinus amblyscalaris (Thurmann), Balanocrinus subteres (Münster), B. pentagonalis (Goldfuss), Isocrinida indet., comatulid Hrabalicrinus zitti gen. et sp. nov., cyrtocrinids Lonchocrinus sp., Cyrtocrinus cf. nutans (Goldfuss), Pilocrinus moussoni (Desor), Tetracrinus moniliformis Münster, Cyrtocrinida indet., and millericrinids (Millericrinida indet.). Crinoids already mentioned from the Jurassic strata of the Czech Republic are subsequently revised. Apart from the isolated remains of Isocrinida, Millericrinida, and Thiolliericrinida, the presence of any other taxon reported from this area should be treated with extreme caution. A rare example of non-regenerative columnal healing (the so-called callus) in I. amblyscalaris is also described. Based on sedimentology and microfacies, the Jurassic limestones were deposited in various palaeoenvironments of upper carbonate platform and shelf lagoon (0 to > 50 m palaeodepth).     

The origin of echinoid shell beds in siliciclastic shelf environments: three examples from the Miocene of Sardinia,Italy

Three mass accumulations of sea urchins from the Miocene of Sardinia show a number of taphonomic features which set them apart from previously described echinoid assemblages from the Cenozoic in which they represent: (1) monotypic assemblages; (2) include very well‐preserved remains of either regular or spatangoid echinoids; and (3) originate in deeper water environments. These accumulations are compared using a detailed sedimentological and taphonomic analysis including preservational fabrics, taphonomic signatures, size frequency distributions, density of occurrences and preferred orientations. The possible role of gregarious behaviour contributing to mass occurrences and the specific sedimentary events leading to the excellent preservation are discussed. The interpreted depositional environment of all three deposits is that of a storm‐dominated, siliciclastic shelf environment. A phymosomatid assemblage represents rapid burial through obrution of a highly dense, freshly dead community. A Brissopsis‐dominated spatangoid assemblage represents a mixed accumulation of parautochthonous and transported skeletons. The third assemblage consisting of regular echinoid spines and rare tests represents a composite tempestite. Differences in the depositional environments are related to their position along onshore–offshore gradient with the first two beds originated in a deeper setting than that of the spine accumulation. This study shows that the preservation of assemblages containing complete regular echinoids and spatangoids is higher in deeper water settings than in shallow water environments.     

Echinoid test damage by a stingray predator

Stingrays (Order: Myliobatiformes) are well‐known predators that feed on a variety of marine species including crustaceans, teleosts, molluscs and annelids. Here, a predatory attack by a stingray on the large spatangoid echinoid Meoma ventricosa (Lamarck) is reported from the shallow‐water carbonate platform of San Salvador, Bahamas. A single stingray was observed feeding on a large adult specimen of M. ventricosa near Sand Dollar Beach, San Salvador Island. The partially crushed and partially eviscerated test of the attacked echinoid was collected from the feeding site. In addition, a crushed test of a freshly killed, smaller irregular clypeasteroid echinoid Leodia sexiesperforata (Leske) was also recovered at the same site. In both cases, the test was broken across the test plates and not along plate boundaries. Meoma ventricosa showed major plate loss on the oral side; however, bite traces and spine loss were not observed in the damaged areas. Similarly, L. sexiesperforata showed no evidence of spine removal; however, bite marks were identifiable. The observed test damage in both taxa is morphologically distinct and could be potentially identifiable in the fossil record.     

Peltoceratinae (Ammonoidea) from the upper Callovian–lower Oxfordian of Kutch,India: Systematic revision and biostratigraphic implication

The ammonoid subfamily Peltoceratinae is a well-constrained group restricted to the Middle Upper Jurassic; it had a near global distribution. In Kutch (India), they were the most important marine macroinvertebrate group during the upper Callovian–lower Oxfordian. Previous reports suffered from excessive splitting due to failure to account for the large intraspecific variability and sexual dimorphism of these ammonoids. The present systematic revision, based on numerous specimens including many near-complete adult specimens, enables to recognize and describe three species of the genus Peltoceras (P. athleta, P. ponderosum, and P. kumagunense) and two species of the genus Peltoceratoides (P. semirugosus and P. propinquus), as well as identifying their macroconchs and microconchs. We rediscovered Peltoceras athleta, which was previously described by Waagen (1875) but often overlooked by subsequent workers. We were able to identify several morphs within this species that are stunningly similar with those previously reported from France. The presence of P. athleta sensu stricto prompted us to reinstate the Athleta Zone in Kutch, improving intercontinental biostratigraphic correlation. Also, it highlights that the genus Peltoceras has a different biostratigraphic longevity in Kutch: in Europe it is restricted to the upper Callovian, whereas in India it also occurs in the lower Oxfordian.     

New Ichnospecies of Cardioichnus from the Miocene of the Guadalquivir Basin,Huelva, Spain

A new ichnospecies of Cardioichnus, Cardioichnus reniformis isp. nov., is documented and described from marine strata of the Late Miocene of southwestern Spain. It is a bilobate resting trace with a heart‐shaped outline and is related to the work of a wedge‐shaped irregular echinoid of the spatangoid group.     

Bichordites monastiriensis from the Pleistocene of southeast Jamaica

Bichordites monastiriensis Plaziat and Mahmoudi 1988, produced by burrowing spatangoid echinoids, is figured and described from shallow marine siliciclastic strata of the Pleistocene Old Pera Beds (Coastal Group) of southeast Jamaica. Its occurrence there represents only the second formal recording of this monospecific ichno‐taxon and extends its geographic range from the Mediterranean region to the Caribbean. The spatangoid species that produced this trace fossil is unknown, but, unlike examples from other areas, it is unlikely to be Echinocardium.     

Répartition stratigraphique et biozones des kystes de dinoflagellés au passage Jurassique moyen–Jurassique supérieur (Bathonien supérieur–Oxfordien inférieur) dans le Bassin de Guercif,Maroc nord-oriental

Despite the stratigraphical significance of dinoflagellate cysts as reliable markers for correlating and dating Jurassic–Cretaceous strata, investigations into this palynomorph group in the southern Tethyan Realm, specifically northwest Africa, are sparse and somewhat parochial. Most research on Jurassic dinoflagellate cysts is focussed on European depocentres in the Boreal and Sub-Boreal realms. This study is on biostratigraphical data from two petroleum boreholes (MSD1 and KDH1) drilled during 1985 in the Guercif Basin, northeast Morocco by ONAREP (Office National de Recherche et d’Exploitation Pétrolier), now ONHYM (Office National des Hydrocarbures et des Mines). These boreholes penetrated a thick siliciclastic succession, attributed to the Middle and Upper Jurassic, below Miocene marls. Over sixty dinoflagellate cyst taxa were identified. Four dinoflagellate cyst biozones, named GI to GIV, are established for the late Bathonian to early Oxfordian interval. These biozones are defined on the basis of the first appearance datum (FAD) and/or the last appearance datum (LAD) of some biomarker taxa which have wide geographical distributions. These are: Ctenidodinium combazii and Ctenidodinium sellwoodii for the GI Biozone (late Bathonian–early Callovian); Ctenidodinium continuum and Meiourogonyaulax caytonensis for the GII Biozone (middle Callovian); Gonyaulacysta centriconnata and Wanaea thysanota for the GIII Biozone (late Callovian–earliest Oxfordian); and Liesbergia liesbergensis and Systematophora penicillata for the GIV Biozone (early Oxfordian). These biozones are correlated to those already established for the respective intervals in other palaeogeographic regions, such as the Boreal, Sub-boreal and Tethyan realms.     

Cyrtocrinids (Echinodermata,Crinoidea) from Upper Jurassic Štramberk‐type limestones in southern Poland

Abstract: A systematic account of highly diverse cyrtocrinid faunules from Upper Jurassic strata of ?tramberk type (Oxfordian–Tithonian) in southern Poland (Polish Carpathians) is presented. Fourteen taxa (Phyllocrinus malbosianus, Ph. stellaris, Ph. sp., Psalidocrinus armatus, Sclerocrinus compressus, S. polonicus sp. nov., Hemicrinus aff. kabanovi, Ancepsicrinus parvus gen. et sp. nov., Tetracrinus baumilleri sp. nov., Eugeniacrinites alexandrowiczi, E. cf. moravicus, E. sp., Eudesicrinus gluchowskii sp. nov. and Hemibrachiocrinus tithonicus sp. nov. are described and illustrated. Representatives of the genus Eudesicrinus, previously recorded only from the Lower Jurassic, are here shown to extend into the uppermost Jurassic. Other cyrtocrinids considered are common in Jurassic/Cretaceous strata across Europe. In the present faunules, isocrinid (Isocrinida), comatulid (Comatulida) and roveacrinid (Roveacrinida sensu Rasmussen, inclusive of Saccocoma) crinoids are associated.     

Corals,molluscs, and paleogeography of late Jurassic strata of the Cerro Pozo Serna,Sonora, Mexico

Jurassic strata of late Oxfordian to early Kimmeridgian age are reported from the Cerro Pozo Serna, northwest-central Sonora, Mexico. The presence of these marine strata greatly alters previous paleogeographic reconstructions of the Tethyan embayment in this area. Approximately 60% of the Sonorian molluscan fauna has been previously reported from Jurassic horizons in the Gulf of Mexico and west Texas regions. The remainder of the molluscs have previously been reported from either the western interior United States or Canada. The Pozo Serna faunas seem to be zoogeographically transitional between communities present in the southern North American Tethyan realm and regions within the central North American Tethyan realm. Additionally, the Pozo Serna fauna greatly resembles contemporaneous communities reported from the Jurassic of Morocco. Corals from the Sonorian Jurassic prove to be, in large part, conspecific with well-known central European and Algerian reef-building species. The scleractinian Lepidophylliopsis gen. nov. and Macgeopsis sonorensis sp. nov. are described from central Sonora.     

Diversity of Cenozoic marsupiate echinoids as an environmental indicator

McNamara, K.J. 1994 10 15: Diversity of Cenozoic marsupiate echinoids as an environmental indicator. Marsupiate echinoids are today largely confined to the seas around Antarctica. Consequently, it has often been inferred that the presence of marsupiate echinoids in the fossil record is indicative of the former existence of low oceanic temperatures. In this study the distribution of marsupiate echinoids through the Cenozoic succession of southern Australia is compared with palaeo-temperature data to test this assumption. The analysis reveals that there is no positive correlation between high marsupiate echinoid diversity during the Cenozoic and low oceanic temperatures. An alternative hypothesis, based on life-history strategies, is investigated. This reveals that marsupiate echinoids show many characteristics typical of organisms with slow growth, long life spans and production of few, large offspring. It is suggested that the northward migration of Australia during the Cenozoic from an original high-latitude location in the early Cenozoic was accompanied by an increase in environmental instability in the southern Australian region in the late Cenozoic. This led to a consequent decrease in marsupiate echinoid diversity. During the Pliocene these direct brooding echinoids were replaced by non-brooders with pelagic lecithotrophic larvae, which dominate the southern coastal echinoid fauna of Australia today. The environmental stability experienced in southern Australia in the early Cenozoic persisted throughout the Cenozoic in the Antarctic region, particularly with regard to predictability of nutrient supply. The result has been the dominance of marsupiate echinoids in that region today. Temporal changes in the diversity of marsupiate echinoids in southern Australia therefore supports the view that their spatial and temporal distribution may be more closely correlated with aspects of their life-history strategy and environmental stability than with low temperature. Echinoidea, evolution, diversity, life-history strategy, Cenozoic.     

A new phosphatic‐shelled cirripede (Crustacea,Thoracica) from the Lower Jurassic (Toarcian) of Germany – the oldest epiplanktonic barnacle

A new genus and species of phosphatic‐shelled eolepadid barnacle from the Posidonia Shale (Toarcian, falciferum Zone) of Zell u. Aichelberg, southern Germany, is described as Toarcolepas mutans gen. et sp. nov. Numerous disarticulated individuals, associated with fossil wood, are present in a piece of concretionary limestone, and these are interpreted as having lived epiplanktonically attached to driftwood. The taxonomy of the Late Triassic – Early Cretaceous family Eolepadidae is reviewed, and two further species (T. gaveyi (Withers, 1920) and T. lotharingica (Méchin, 1901)) are referred to Toarcolepas. The chemistry of valve composition of the Carboniferous Praelepas and Triassic–Jurassic eolepadid cirripedes is investigated using X‐ray dispersive analysis, and the ubiquitous presence of abundant phosphorus is taken as evidence that these taxa had a primary phosphatic composition, now preserved as francolite. A significant change in shell chemistry from phosphate to calcium carbonate took place during the evolution of the Thoracica, during the Late Triassic or Early Jurassic. The driving force behind this change may have been related to the reduced predation pressure associated with acquisition of an epiplanktonic mode of life. Calcite is softer, but energetically cheaper to deposit than phosphate mineral phases.     

Shallow Traces (Pits) in the Test of the Irregular Echinoid Echinocorys scutata Leske from the Chalk (Upper Cretaceous) of the United Kingdom

Specimens of epifaunal irregular echinoids in the Upper Cretaceous of northern Europe have been reported with patterns of circular, nonpenetrative parabolic pits, Oichnus paraboloides (Bromley), in the apical region. Specimens of Echinocorys scutata Leske from the Chalk at two sites in southeast England were commonly penetrated by this trace, generated by an indeterminate pit-forming organism. Pits commonly surround the apical system and, less commonly, occur within it; they occur preferentially anteriorly. Pits occur within plates, not along margins or sutures. Crosscutting of pits indicates that multiple spatfalls probably occurred. The host echinoid added new test plates adjacent to the apical system; thus, plates bearing O. paraboloides were moved abapically. The reduction in number of pits away from the apex, including those with echinoid tubercles reestablished on the base, indicates that, following death of an infester, the echinoid “reclaimed” and infilled them with calcite. The pit-former was most probably an unmineralized invertebrate that used E. scutata as a domicile which provided good access to food-rich currents for suspension feeding. Although the systematic position of the pit-former is unknown, similar infestations are known from other Upper Cretaceous echinoids and Mississippian crinoids.