Living on an island: characterization of the encrusting fauna of large pectinid bivalves from the Lower Cretaceous of the Neuquén Basin,west‐central Argentina

Exposed mollusc shells may act as benthic islands in soft bottoms, and the analysis of their encrusting faunas provides unique palaeoecological information. In the late Valanginian of the Agrio Formation (Neuquén Basin, west‐central Argentina), the large pectinid Prohinnites acted as a benthic island on soft substrates. Inequivalved Prohinnites adults with small, smooth cementing scars on the right valve suggest that a free reclining life habit followed the epibyssate juvenile and cementing phases. The encrusting fauna on Prohinnites was studied taxonomically and palaeoecologically by means of a quantitative approach. Over 90% of 123 valves presented encrusters. Encrustation was equally common in both valves. Internal encrustation was rare. The left umbonal region was less encrusted probably due to sediment accumulation or early colonization by soft‐bodied taxa. The fauna was composed of 14 encrusting taxa, including oysters, serpulids, sabellids and cyclostome bryozoans. Oysters exceeded 50% of the total abundance, but serpulids and bryozoans were more diverse. Serpulids and particularly oysters showed a gregarious life habit. Few interactions took place among encrusters and most were post‐mortem, involving the overgrowth of already dead oysters. The oysters were early settlers that took advantage of their gregarious behaviour to rapidly cover available hard surfaces. However, they were unable to exclude bryozoans and polychaetes, which settled on the pectinid's valves regardless of the presence of oysters. The studied fauna corresponds to a climax community that was structured by larval abundance rather than by competitive interactions; oysters settled first and replenished themselves while polychaetes and bryozoans settled over or alongside them     

Palaeoecological analysis of a sclerobiont fauna on a single basibiont across the Valanginian of the Neuquén Basin,west‐central Argentina

Sclerobiont communities have proven their environmental and taphonomic value in extant and past settings; studies are beginning to quantify and evaluate their changes across time. Through the Valanginian of the Neuquén Basin, Argentina, trigonioids of the genus Steinmanella present an almost continuous record. Materials were collected from shales and shell beds across second‐order systems tracts (LST, TST and HST). Their sclerobiont communities were characterized and compared across systems tracts and facies. In addition, a link between them and coeval oyster mass occurrences (OMOs) was investigated, since studies on other local sclerobiont communities have consistently shown pronounced oyster dominance. Eleven sclerobiont taxa were found on Steinmanella (bryozoans, bivalves, foraminifers and polychaetes), of which oysters are usually strongly dominant but for the TST, in which their dominance is diminished. The sclerobiont community has a simple structure and interspecific relationships and competition for space seem to have had little importance. Few differences in richness, taxonomy and other parameters were found between facies. Across systems tracts, features of the sclerobiont communities are differentiated mostly from swings in relative abundance of taxa. The lowered oyster dominance in the TST may reflect a source–sink dynamics between OMOs (sources) and Steinmanella as sinks in shallower, soft substrate settings. Likely the sea‐level rise temporally drowned the oyster carbonate factory, resulting in decreased larvae emigration to the sinks and lowered oyster dominance during the TST.     

Microtaphonomy of bioclasts and paleoecology of microencrusters from Upper Jurassic spongiolithic limestones (External Prebetic,Southern Spain)

Microencrusters and microtaphonomic features of the Oxfordian spongiolithic limestones of the External Prebetic were studied using thin-section analysis. The spongiolithic limestone is a bioclastic-rich packstone with common echinoderm, mollusc and brachiopod remains. The bioclasts show a high fragmentation index and frequent microborings. The encrustation index (E _i) is higher for fragments of serpulids, ammonoids and bivalves, and increases with the initial grain-size of bioclasts. The main microencrusters consist of benthic microbial communities (BMC) and nubeculariids, as well as subordinate calcareous and siliceous agglutinated foraminifera, serpulids and bryozoans. BMC are usually the first colonizers, and encrusting foraminifera mainly appear on bioclasts larger than 2 mm. BMC dominate in well-developed encrustations on upward facing surfaces of larger bioclasts that are also colonized by foraminifera (nubeculariids and Subdelloidina). Bullopora, serpulids and bryozoans are more common on lower surfaces. The fact that the values of encrustation index, encrustation thickness and diversity of the microencrusters increase with the size of bioclasts is related to a higher stability and exposure time of the available bioclastic substrate. The microencruster distribution on upper and lower surfaces of large bioclasts may be related to photic control, space competition and/or predation avoidance.     

Taphonomy and palaeoecology of Hauterivian–Barremian nerineoid shell beds from the Neuquén Basin,west‐central Argentina

Nerineoid shell beds are described for the first time from Lower Cretaceous deposits of southern South America. These come from carbonates near the top of the Agrio Formation in southern Mendoza Province, west‐central Argentina. To envisage the origin of the nerineoid shell beds, a taphonomic study was carried out, which indicated that these represent within‐habitat time‐averaged, primary sedimentological concentrations with a secondary biogenic imprint related to a relatively high local production of nerineoid shells. The associated palaeoenvironments were studied through a facies analysis of the carbonate succession including the shell beds. The carbonates were deposited in a homoclinal ramp system and depict a shallowing upward trend from mid to inner ramp. The individuals lived and accumulated in oolitic shoals within the inner ramp, in a shallow, well‐lit, high‐energy setting above fair‐weather wave base. Substrate was oxygenated and loose. The nerineoids are shown to belong to one species of the genus Eunerinea, and through the functional morphology of the shells they are tentatively interpreted as infaunal or semi‐infaunal. It is suggested that the recorded monospecific nerineoid shell beds indicate that the palaeoenvironmental conditions may have been favourable for the development of abundant populations of these gastropods in the northern part of the Neuquén Basin during a short time interval in the Hauterivian–Barremian boundary. This could have been related to a brief warming episode, but other factors may have also been involved. □Argentina, Early Cretaceous, Gastropods, nerineoids, Neuquén Basin, shell beds, taphonomy.     

Late Jurassic ichthyosaurs from the Neuquén Basin,Argentina

Late Jurassic ichthyosaurs are well represented in the Tithonian of the Neuquén Basin, in northwestern Patagonia, Argentina. Most of the ichthyosaur material from the Neuquén Basin was originally identified as Ophthalmosaurus. Recently, the new ichthyosaur genus Caypullisaurus was described, based on an almost complete mature specimen from Cerro Lotena. Some material previously referred to Ophlhalmosaurus has been referred to the new genus. However, both genera are present in the Tithonian of the Neuquén Basin. The discovery of an articulated forefin in Cajón de Almanza (near Loncopue, Neuquén) confirms the presence of Ophthalmosaurus in the uppermost Tithonian of the Neuquén Basin.     

Morphological trends and environment: a case study on Early Cretaceous Myophorelloidea (Bivalvia,Trigoniida) from Neuquén Basin,Argentina

Trigoniide clams were conspicuous components of bivalve faunas in shallow seas during most of the Mesozoic. Morphological trends in different features of shell shape and ornamentation of three related species of myophorelloids from the Agrio Formation (Early Cretaceous of Neuquén Basin, Argentina) that inhabited environments of different hydrodynamic energy conditions are explored in this study by means of a detailed morphometric analysis. Four hydrodynamic energy categories were established for depositional environments, from most (1), to least (4), energetic. General shell shape was characterized by measurements of height, length and width. New quantitative methods were developed to evaluate the distribution of costae along the marginal carina and the divergence of costae along the flank. Morphometric variables were detrended with respect to size, to avoid ontogenetic variability in further analyses. Relative width of the shell increases as environments become higher in hydrodynamic energy; this unusual trend may be interpreted as an anchoring/stabilizing strategy developed by the lineage in shallow marine environments. Trends in ornamentation correspond to more densely distributed costae in environments of higher hydrodynamic energy, a pattern that improves substrate penetration.     

Ontogeny and evolution within the Myophorellidae (Bivalvia): Paedomorphic trends

The role of heterochronic phenomena in molluscan evolution is insufficiently understood but potentially significant. The aim of this paper is to explore some paedomorphic trends in the evolution of the Myophorellidae (Bivalvia: Trigoniida). Early ontogeny of general shell shape and ornamentation of one species of Steinmanella was analyzed and compared to data obtained for three species of Myophorella: two belonging to the subgenus M. (Promyophorella) (one from the Jurassic and one from the Cretaceous) and one belonging to the Jurassic M. (Myophorella). For general shell shape, a geometric morphometric analysis was performed on lateral views of the shells. Regarding ornamentation, flank costal disposition on the marginal carina, tubercle separation and relative development of the sub-commarginal subset of flank costae were quantified. A qualitative analysis was also performed. A two-trend shell shape development is considered as primitive. The first trend is marked by a relative reduction of the posterior margin together with a relative elongation of the shell. A tangential opisthogyrate growth component characterizes the second trend. There is a transitional stage where both trends interact. Early flank ornamentation is characterized by two or three sub-commarginal costae, continuous through the area, after which oblique costae with fine tubercles start to form. The subgenus M. (Myophorella) evolved by paedomorphic retention of juvenile shell shape and ornamentation, resulting in a large shell with coarse tubercles. Shell morphology in Steinmanella evolved by paedomorphic suppression of the primitive second trend in the development of the shell, resulting in an orthogyrate shell shape, and the retention of juvenile ornamentation (coarse tubercles, more sub-commarginal costae, juvenile rates of costal disposition). The paedomorphic (most likely by deceleration) retention of juvenile shell morphology within the Myophorellidae seems to have been recurrent within the group, resulting in many cases of convergence, and obscuring the phylogenetic relationships among its species.     

Large gryphaeid oysters as habitats for numerous sclerobionts: a case study from the northern Red Sea

The shell of a living specimen of the Indo-Pacific gryphaeid giant oyster Hyotissa hyotis was colonized by numerous encrusting, boring, nestling and baffling taxa which show characteristic distribution patterns. On the upper valve, sponge-induced bioerosion predominates. On the lower valve intergrowth of chamid bivalves and thick encrusting associations—consisting mostly of squamariacean and corallinacean red algae, acervulinid foraminifera, and scleractinian corals—provides numerous microhabitats for nestling arcid and mytilid bivalves as well as for encrusting bryozoans and serpulids. Such differences between exposed and cryptic surfaces are typical for many marine hard substrata and result from the long-term stable position of the oyster on the seafloor. The cryptic habitats support a species assemblage of crustose algae and foraminifera that, on exposed surfaces, would occur in much deeper water.     

An exceptional storm accumulation of nautilids in the Lower Cretaceous of the Neuquén Basin,Argentina

Cichowolski, M., Pazos, P.J., Tunik, M.A. & Aguirre‐Urreta, M.B. 2011: An exceptional storm accumulation of nautilids in the Lower Cretaceous of the Neuquén Basin, Argentina. Lethaia, Vol. 45, pp. 121–138. An exceptional accumulation of nautilid shells of the species Cymatoceras perstriatum (Steuer) is reported and described in the Lower Cretaceous Agrio Formation of the Neuquén Basin (west‐central Argentina). The bed represents a storm deposit in a shallow‐water environment within the mid‐ramp. The evidences of a storm‐related origin of the bed come from the petrographic analysis and taphonomic features of the shells, specially the sedimentary infill pattern. The shells are dispersed in patches within the stratum, without any orientation relative to the bedding plane. It is proposed that the shells were floating after the death of the animals, although most of them have an almost complete living chamber. The presence of some heavily encrusted shells suggests that there is a mixing of specimens with different drift times. A variety of factors related to the origin of such high number of nautilid shells are discussed, including a transgressive stand system tract, the possible existence of a gregarious behaviour, changes in salinity and wind directions. □Lower Cretaceous, Nautilids, Neuquén Basin, shell accumulation, storm deposit.     

Early cretaceous serpulid limestones: Chachao formation,Neuquen basin,Argentina

Summary Colonial serpulids (Sarcinella) are common within some limestones of the lower Chachao Formation (Valanginian) near Malargüe city at the northern border of the Neuquem basin, Argentina. The shallowing-upwards sequence is characterized by low-energetic micritic carbonates. The serpulid limestones mark the transition from a carbonate ramp stage to a platform stage.     

Epizoan encrustation on Marsupites testudinarius – additional argument favoring an epifaunal mode of life of uintacrinoids?

Uintacrinoids (Uintacrinoidea Zittel) are among the best-known Late Cretaceous crinoids, but owing to their unusual morphology and widespread distribution their mode of life has become a subject of much discussion. Of several competing hypotheses, nektonic, pseudoplanktonic, hemipelagic, semi-infaunal and epibenthic lifestyles have been suggested. Recent study synthesizing and extending previous data has shown that these crinoids were epibenthic, holding their arms vertically for feeding. However, evidence supporting a rheophilic epibenthic model over an alternative rheophobic semi-infaunal model is still limited. Here we report epizoans, mostly represented by serpulids and bryozoans, encrusting thecal plates of Marsupites testudinarius from the Lägerdorf in Germany. Although a definitive interpretation whether recorded infestations occurred syn vivo or post mortem is not certain, it is remarkable that all epizoans (or their traces) are attached to the convex side (latera) of well-preserved isolated plates displaying no signs of reworking. Furthermore, a bryozoan colony crossing plate boundaries has been also found on the surface of a sub-articulated theca suggesting that it may have been colonised syn vivo. Recorded epibiotic associations, whether syn vivo or post-mortem, must have developed prior to burial of the specimens, above the surface of sea floor, and thus provide another argument against rheophobic semi-infaunal mode of life of uintacrinoids.     

Mode of life of the Silurian uncoiled gastropod Semitubina sakoi n. sp. from Japan

Semitubina sakoi n. sp. from the late Silurian of Japan represents the second species of this genus and also the first record of a Silurian gastropod in Japan. The gastropod shells occur in a thin mudstone bed and were found to be encrusted exclusively by corallites of ? Favosites sp. These corallites reveal that encrustation proceeded as the gastropod shells grew. The ecological relationship between the two organisms is considered to be symbiotic. This mode of life allowed the coral to live on a muddy substrate because clear sea water passed over the colony as the gastropod moved along. The gastropod benefited from this relationship by being protected from shell-boring or shell-crushing predators by the encrusting corallite. In Semitubina sakoi the body whorl is separated from the penultimate one by a considerable gap in a later growth stage and S. sakoi has been cited as one of the uncoiled gastropods. The uncoiling of this gastropod results primarily from rapidly increasing whorl translation rate in the latest growth stage. Taking the symbiotic relationship with ? Favosites sp. into consideration, a deposit feeding or benthic scavenging mode of life is suggested for this gastropod.     

Upper Eocene larger foraminiferal–coralline algal facies from the Klokova Mountain (southern continental Greece)

Within the Gavrovo–Tripolitza area (southern continental Greece), marine carbonate platforms existed from the Late Triassic to the Late Eocene. The Middle–Upper Eocene marine shallow-water carbonates of the Klokova Mountain represent remnants of the large volumes of sediment that were produced on a middle ramp sedimentary system which culminated in the Lower Oligocene terrigenous deposits. Facies analysis of Bartonian–Priabonian shallow-water carbonate successions and the integration with palaeoecological analysis are used to produce a detailed palaeoenvironmental model. In the proximal middle ramp, porcelaneous foraminiferal packstone facies is characterised by larger foraminifera such as Praturlonella and Spirolina. These forms thrived in a shallow-water setting with low turbidity, high-light intensity and low-substrate stability. The foraminiferal packstone facies, the thin coralline wacke–packstone facies and the rhodolith packstone facies deposited approximately in the same depth range adjacent to one another in the middle-ramp. Nummulitids (Nummulites, Assilina, Pellatispira, Heterostegina and Spiroclypeus) increase in abundance in the middle to distal mid-ramp together with the orthophragminids. Coralline algae, represented by six genera, are present in all facies. Rhodoliths occur in all facies but they show different shapes and growth forms. They develop laminar sub-ellipsoidal shapes in higher turbulence conditions on mobile sand substrates (foraminiferal packstones and rhodolith rudstones), whilst sub-discoidal shapes often bound by thin encrusting coralline plants in lower hydrodynamic settings. The distinctive characteristics of the palaeoecological middle-ramp gradient are an increase in dominance of melobesioids, a thinning of the encrusting coralline plants and a flattening of the larger benthic foraminiferal shells.     

Decapod Crustacea from the Agrio Formation (Lower Cretaceous) of the Neuquén Basin,Argentina

Abstract: Two species of decapod crustacean are recorded from the Agua de la Mula Member of the Agrio Formation (Upper Hauterivian – Lower Barremian) of the Neuquén Basin of west‐central Argentina, namely Astacodes falcifer Bell and a new species of Palaeohomarus, P. pacificus. The preservation of the specimens is exceptional, some showing delicate compound eyes and a stridulatory apparatus, features rarely found in fossil forms. Many specimens are preserved articulated inside calcareous nodules, within dark‐grey shales. The lobster‐bearing sediments accumulated in a low‐energy marine environment and diagenetic mineralization occurred very rapidly, prior to significant decay, thus allowing exceptional preservation of specimens. Palaeohomarus was a rare genus in the Cretaceous with a palaeogeographic distribution restricted to the Mediterranean Tethys, the eastern USA and Madagascar, while Astacodes falcifer has been recorded only from Speeton (eastern England) and Neuquén.     

Multi‐proxy analyses of Late Cretaceous coprolites from Germany

A total of 462 coprolites from three localities exposing Upper Cretaceous deposits in the Münster Basin, northwestern Germany, have been subjected to an array of analytical techniques, with the aim of elucidating ancient trophic structures and predator–prey interactions. The phosphatic composition, frequent bone inclusions, size and morphology collectively suggest that most, if not all, coprolites were produced by carnivorous (predatory or scavenging) vertebrates. The bone inclusions further indicate that the coprolite producers preyed principally upon fish. Putative host animals include bony fish, sharks and marine reptiles – all of which have been previously recorded from the Münster Basin. The presence of borings and other traces on several coprolites implies handling by coprophagous organisms. Remains of epibionts are also common, most of which have been identified as the encrusting bivalve Atreta. Palynological analyses of both the coprolites and host rocks reveal a sparse assemblage dominated by typical Late Cretaceous dinoflagellates, and with sub‐ordinate fern spores, conifer pollen grains and angiosperm pollen grains. The dinoflagellate key taxon Exochosphaeridium cenomaniense corroborates a Cenomanian age for the Plenus Marl, from which most studied coprolites derive. The findings of this study highlight the potential of a multi‐proxy approach when it comes to unravelling the origin, composition and importance of coprolites in palaeoecosystem analyses.     

Recent abyssal benthic foraminiferal biofacies of the eastern equatorial Indian Ocean

Study of Recent abyssal benthic foraminifera from core-top samples in the eastern equatorial Indian Ocean has identified distinctive faunas whose distribution patterns reflect the major hydrographic features of the region. Above 3800 m, Indian Deep Water (IDW) is characterized by a diverse and evenly-distributed biofacies to whichGlobocassidulina subglobosa, Pyrgo spp.,Uvigerina peregrina, andEggerella bradyi are the major contributors.Nuttalides umbonifera andEpistominella exigua are associated with Indian Bottom Water (IBW) below 3800 m. Within the IBW fauna,N. umbonifera andE. exigua are characteristic of two biofacies with independent distribution patterns.Nuttalides umbonifera systematically increases in abundance with increasing water depth. TheE. exigua biofacies reaches its greatest abundance in sediments on the eastern flank of the Ninetyeast Ridge and in the Wharton-Cocos Basin. The hydrographic transition between IDW and IBW coincides with the level of transition from waters supersaturated to waters undersaturated with respect to calcite and with the depth of the lysocline. Carbonate saturation levels, possibly combined with the effects of selective dissolution on the benthic foraminiferal populations, best explain the change in faunas across the IDW/IBW boundary and the bathymetric distribution pattern ofN. umbonifera. The distribution of theE. exigua fauna cannot be explained with this model.Epistominella exigua is associated with the colder, more oxygenated IBW of the Wharton-Cocos Basin. The distribution of this biofacies on the eastern flank of the Ninetyeast Ridge agrees well with the calculated bathymetric position of the northward flowing deep boundary current which aerates the eastern basins of the Indian Ocean.     

The Early Cretaceous of North-East Greenland: A crossroads of belemnite migration

Recent field work in Lower Cretaceous successions of Traill Ø and Wollaston Forland, North-East Greenland, have resulted in c. 2350 belemnite guards collected bed-by-bed from the upper Ryazanian – Hauterivian. The most common belemnite genera observed, Acroteuthis, Pachyteuthis, and Cylindroteuthis are of boreal-arctic affinities and closely related to NW European and Siberian faunas. Other taxa, including Hibolithes (common), Pseudobelus (relatively common) and Duvalia (rare), show faunal links to both NW European and Mediterranean faunas. This paper describes and discusses these findings in their taxonomic, biostratigraphic, palaeobiogeographic, palaeoecologic and palaeoceanographic context. In particular, the occurrence of Pseudobelus which is common in the circum Mediterranean area, is remarkable since it is the first observation of this Tethyan genus in the entire Boreal Realm. The palaeoecological interpretation of these observations result in the recognition of four different palaeobiogeographic belemnite assemblages for the Boreal Realm: 1) North-East Greenland, 2) Spitsbergen, 3) NW Europe and 4) Siberia. In contrast to the other assemblages, the belemnite faunas of North-East Greenland consist of a) Boreal-Arctic elements, b) Boreal-European taxa, c) endemic belemnites of Tethyan ancestry, and d) Tethyan species. These findings make North-East Greenland part of an immigration route from the Tethyan Realm via the north Atlantic to the high Boreal. This allowed Tethyan species, which are otherwise unknown from the Boreal Realm, to reach North-East Greenland. The occurrence of the Tethyan genus Pseudobelus in North-East Greenland also supports the interpretation of this taxon as a hemipelagic dweller, capable of crossing major distances. The belemnite patterns further suggest the existence of a proto Gulf-stream, documenting a south-to-north flow of warm surface waters as far north as Greenland already in the earliest Cretaceous (Valanginian). This has substantial implications for the interpretation of Early Cretaceous climate and oceanic current systems, as well as for the palaeobiology of belemnites.     

Mosasaur predation on a nautiloid from the Maastrichtian Pierre Shale,Central Colorado,Western Interior Basin,United States

Mosasaurs were common predators on the ammonites that inhabited the upper water column of the Late Cretaceous Western Interior epicontinental seaway of North America. Mosasaurs developed predictable behaviour patterns for feeding on ammonite prey. There are no previous reports of mosasaur predation on the much less common Cretaceous nautiloids, possibly because of the prey's predominantly deep, epibenthic habitat, as deduced from modern Nautilus life habits. A single specimen of the highly inflated nautiloid, Eutrephoceras dekayi (Conrad), prey to a small adult mosasaur, likely Platycarpus, Prognathodon or Mosasaurus, is reported herein from the Pierre Shale of Colorado in the Early Maastrichtian biozone of Baculitesgrandis transitional to the biozone of B. clinolobatus. The nautiloid was killed in the same manner as described previously for discoid ammonites (Placenticeras, Sphenodiscus) from coeval strata in the USA and Canada.     

Rogerella isp. Infesting the Pore Pairs of Hemipneustes striatoradiatus (Leske) (Echinoidea: Upper Cretaceous,Belgium)

An otherwise well-preserved test of the holasteroid echinoid Hemipneustes striatoradiatus (Leske) from the Emael Member, Maastricht Formation (Maastrichtian, Upper Cretaceous) of Belgium, is infested by encrusting bivalves and foraminiferans and the boring Rogerella isp. In this specimen, Rogerella preferentially infested and modified the ambulacral pore pairs of the echinoid close to the apex. This was not a commensal association. The echinoid test shows no growth deformations in response to this invasion; pore pairs are locally strongly infested; and encrusting invertebrates testify to the long post-mortem residence time of the test on the sea floor. Rather, the pore pairs of the dead echinoid were crannies attractive to settling larvae of acrothoracian barnacles, the producers of Rogerella.