Late Cambrian hard substrate communities from Montana/Wyoming: the oldest known hardground encrusters

Hardground surfaces from the Late Cambrian Snowy Range Formation in Montana/Wyoming are the oldest known non-reefal hard substrates exhibiting encrusting fossils. These surfaces range in age from Early Franconian to early Trempealeauan. Hardgrounds were developed on slightly hummocky to planar, truncated surfaces of glauconite-rich, carbonate, flat pebble conglomerates, which were deposited during episodes of storm scouring in shallow subtidal environments of the Montana/Wyoming shelf. Snowy Range hardgrounds are encrusted by a low diversity assemblage of fossils dominated by simple discoidal holdfasts of pelmatozoans, probably crinoids, and including small conical spongiomorph algae? and probable stromatolites. Macroborings (e.g. Trypanites) are notably absent from all hardground surfaces, although sharp-walled, vertical, cylindrical holes (borings?) occur in micrite clasts imbedded in certain flat pebble conglomerates. No evidence of faunal succession or microecologic partitioning of irregular surfaces was observed on these Cambrian hardgrounds.     

Hettangian (Early Jurassic) Dinosaur Tracksites from the Mecsek Mountains,Hungary

A rare example of a North American Jurassic hardground is found in the Carmel Formation of southwestern Utah. The Carmel hard‐ground was formed across a carbonate lagoon from an oolitic shoal seaward to a subtidal shelly facies landward. It has an abundant bivalve fauna consisting of thick layers of encrusters (the oyster Liostrea and the plicatulid Plicatula), borers (the ichnofossil Gastro‐chaenolites with the mytilid Lithophaga often preserved inside), and nestlers (the mytilid Modiolus). A rare soft‐bodied bryozoan (Arach‐nidium) is preserved by bioimmuration in the attachment scars of Liostrea; this is the first bioimmuration recorded from the Jurassic of North America, and the first bioimmuration recorded from a hard‐ground. The phoronid boring Talpina is present in some Liostrea shells; it was apparently excavated after the death of these oysters. The Carmel hardground community does not contain other fossils, such as serpulids, brachiopods, foraminiferans, and skeletal bryo‐zoans, typical of Jurassic hardgrounds elsewhere. It represents a low diversity molluscan community developed in a restricted marine environment.     

Upper Pliocene bivalve shell concentrations from the Lower Chelif basin (NW Algeria): Systematics,sedimentologic and taphonomic framework

Sedimentologic and palaeontological investigation of the Upper Pliocene Slama Formation in the Lower Chelif Basin (NW Algeria) led us to collect important bivalve assemblages for taxonomic and taphonomic purposes. A rather comprehensive inventory list of Upper Pliocene bivalves from northwestern Algeria is now available and consists of 30 species, 17 of which are extinct ones. Four principal taphonomic attributes were analysed: bioerosion, encrustation, fragmentation, and abrasion. Physical and biogenic sedimentary structures are used for palaeoenvironmental interpretations. The taphonomic, sedimentologic and ichnological characteristics of most of the deposits suggests they originated from discontinuous processes of winnowing and bypassing of sediments, probably due to the action of storms in shallow waters, mainly in the shoreface depositional environment. The bivalve assemblage is dominated by disarticulated valves and displays significant taphonomic alteration in the shells. Sclerobionts traces in shells particulary affect the oyster shells. Bioerosion traces are predominately those of clionid sponges (Entobia isp.), polychaetes (Maeandropolydora isp. and Caulostrepsis isp.), bivalves (Gastrochaenolites isp.), and of predatory gastropods (Oichnus isp.). Among the sclerobionts, the identified encrusters were juvenile oyster recruits, barnacles, polychaetes (serpulid tubeworms), bryozoans (Microporella sp. and Acanthodesia sp.), and vermetid gastropods (Petaloconchus intortus).     

Taphonomic windows and molluscan preservation

Recent studies on silicified fossil biotas have suggested that substantial skewing of the molluscan record resulted from early aragonite dissolution in mid-outer carbonate ramp settings. If those rare skeletal lagerstätten are representative, then the quality and completeness of the molluscan record are thrown into doubt. Yet database studies suggest that the bivalve fossil record is actually relatively complete. If so, then biodiversity must be captured by other processes that preserved shells vulnerable to early dissolution, and which operated on a relatively high frequency, i.e., less than the species duration for bivalves.Storm beds, shell plasters and submarine hardgrounds are identified as fossil deposits that can preserve the labile aragonitic component of the fauna and thus represent potential taphonomic windows. Many storm event beds include rich accumulations of shelly benthos. Differences between storm bed faunas and those of the background facies could reflect transportation effects. However, some storm bed assemblages are rich in originally aragonitic infaunal bivalves that are not represented in background facies or more proximal shelf equivalents, and here rapid burial and removal of organic matter by winnowing may be the keys to aragonite shell preservation. Despite Palaeozoic to Cenozoic changes in the thickness and frequency of shell beds that reflect the predominant bioclast producers, shallow infaunas are commonly concentrated together with epifauna in such deposits.Some low energy, organic-rich mud-dominated settings are associated with preservation of aragonitic molluscs. Infaunal bivalves are a prominent component of shell plasters or pavements in such settings, linked to episodic bottom water anoxia. Decaying algal blooms drew the redox boundary up above the sediment–water interface, and brought populations of infaunal bivalves to the surface where they died. Isolated from the oxic taphonomically active zone, the shells were not dissolved and were buried as thin shell layers. In similar settings, aragonitic shells were preserved as moulds through early pyritisation, or even through preservation of original shell aragonite.In oxic environments, bioturbational reworking of surface sediment destroyed moulds of aragonitic shells after early dissolution. In some hardgrounds, these delicate moulds were preserved due to synsedimentary cementation, probably using carbonate released by aragonite dissolution. The examples included here come from both intervals of “calcite” and “aragonite” seas, and it is not possible to assess whether the saturation state (with respect to aragonite) of the ambient sea water played a role in the selective removal of aragonitic shells.While taphonomic windows may have captured the diversity of individual groups, it is clear from quantitative data involving skeletal lagerstätten that the scale of loss from early aragonite dissolution has drastically altered the trophic composition of some fossil assemblages commonly used as the basis for reconstructions of past communities.     

Middle Jurassic (Bathonian) encrusted oncoids from the Polish Jura,southern Poland

Oncoids from two localities (Ogrodzieniec and Blanowice) of the Polish Jura, southern Poland, have been investigated with respect to their genesis and paleoecology. These oncoids occur within Middle Jurassic (Bathonian) deposits. Those from Ogrodzieniec are large, elliptical, and embedded within a presumably condensed carbonate bed. Those from Blanowice, on the contrary, are significantly smaller, irregular to box-like in shape, and occur within the ore-bearing clays. The oncoids from both localities consist of a distinct carbonate core and laminated cortex that is significantly thicker and better preserved in the Ogrodzieniec oncoids. SEM and optical microscopic investigation of the oncoid cortices revealed the presence of carbonate and silicate layers with web-like structures similar to those occurring in recent cyanobacterial microbialites. Thus, the oncoid cortices investigated may have formed in a photic zone environment with the aid of coccoid and filamentous cyanobacteria. Oxic conditions prevailed during oncoid cortex formation within the siliciclastic setting, which is manifested by low total organic carbon content, high pristane/phytane (Pr/Ph) ratio, and significant predomination of the C₃₁ homohopanes. On the cortices’ surfaces, as well as between particular laminae, various encrusting organisms have been found. The encrusters, dominated by serpulids and bryozoans, are cryptic species that inhabited the undersides and recesses of the oncoids. Their presence on both the upper and lower surfaces of the oncoids indicates that the oncoids were episodically overturned on the seafloor. The much better developed cortex lamination and much higher diversity and abundance of encrusters in the Ogrodzieniec oncoids may point to better trophic conditions prevailing in a shallower marine environment characterized by transparent waters, as opposed to a deeper siliciclastic environment with less transparent waters and probably worse trophic conditions prevailing during formation of the Blanowice oncoids.     

The development of an Early Ordovician hard ground community in response to rapid sea-floor calcite precipitation

The Kanosh Shale (Upper Arenig, Lower Ordovician) of west-central Utah. USA. contains abundant carbonate hardgrounds and one of the earliest diverse hardground communities. The hardgrounds were formed through a combination of processes including the development of early digenetic nodules in clay sediments which were exhumed and concentrated as lags by storms. These cobble deposits. together with plentiful biogenic metrical. were cemented by inorganically precipitated calcite on the sea floor. forming intraformational conglomerate hardgrounds. Echinoderms may have -played a critical role in the development of hardground faunas since their disarticulated calcite ossicles were rapidly cemented by syntaxial overgrowths. forming additional cobbles and hardgrounds. The echinoderms thus may have taphonomically facilitated the development of some of the hard substrates they required. A significant portion of the hardground cements may have been derived from the early dissolution of aragonitic mollusk shells. Kanosh hardground species include the earliest bryozoans recorded on hardgrounds and large numbers of stemmed echinoderms. primarily rhipidocystid cocrinoids. Bryozoans and echinoderms covered nearly equal areas of the hardground surfaces. and there was a distinct polarization between species which preferred the upper. exposed portions of the hardgrounds and others which were most common on undercut. overhang surfaces. The Kanosh Shale hardground fossils combine elements of Late Cambrian assemblages and Middle Ordovician faunas, thus confirming predicted trends in hardground community evolution. especially the replacement of cocrinoids by bryozoans and. to a lesser extent, by other stemmed echinoderms, especially crinoids. The Kanosh community marks the transition from the Cambrian Fauna to The Paleozoic Fauna in The hardground ecosystem. *Carbonate hardgrounds, aragonite dissolution, calcite cement, Echinodermara, Trepostomata, Nicholsonclla. Dianulites. Porifpra. taphonomic facilitation, Utah. Pogonip Group, Kanosh Shale. Ordovician.     

Borings in mobile clasts from Eocene conglomerates of Northern Dalmatia (Coastal Dinarides, Croatia)

Summary Bored clasts occur in Eocene conglomerates deposited in the upper shoreface and beachface settings of the Dinaric foreland basin. The trace fossil assemblage consists ofGastrochaenolites, Trypanites, and possibly some other ichnotaxa and may be compared to theTrypanites Ichnofacies. The preservation characteristics of the borings reflect many stages of colonisation/boring and abrasion. The removal of shells of the boring bivalves, the different depths of the abrasional truncation of borings, and the predominant preservation of the largest excavations (Gastrochaenolites) in the ichnocoenosis are related to repeated phases of abrasion, caused by the mobility of clasts. Coastal gravel is a specific variant of hard substrates, whose mobility controls the colonisation of borers, the type of assemblage and its preservation potential.     

Paleoecology of a Pennsylvanian encrusting colonial rugose coral in south Guizhou, China

The well-preserved Pennsylvanian encrusting colonial rugose coral Ivanovia is widespread and easily observed in south Guizhou, China. There are three common types of hard substrate encrusted by Ivanovia: in situ carbonate hardgrounds, carbonate hardground clasts, and calcareous bioclasts. Thin, spreading sheets are the most common growth form of Ivanovia in the study region. Ivanovia employed peripheral, medial and mixed growth strategies to occupy a sufficient living space on the substrate. It favored a shallow, warm, and clear marine environment within the photic zone and had a high tolerance of water movement. Ivanovia was generally smothered and covered by mud in the studied area. The Ivanovia fossil communities in south Guizhou are characterized by a low species diversity conforming to the typical evolutionary pattern of hard substrate marine communities in the Carboniferous.     

Bryozoans as taphonomic engineers,with examples from the Upper Ordovician (Katian) of Midwestern North America

A combination of encrusting calcitic bryozoans and early seafloor dissolution of aragonitic shells recorded in the Cincinnatian Series of the upper Midwest of North America allowed the preservation of abundant moulds of mollusc fossils bioimmured beneath the attachment surfaces of the bryozoans. We here call this preservational process ‘bryoimmuration’, defined as a bryozoan‐mediated subset of bioimmuration. The bryozoans moulded very fine details of the mollusc shells, usually with more accuracy than inorganic sediment moulds. Most of the bryozoans are heterotrypid trepostomes with robust low‐Mg calcite skeletons. The molluscs are primarily bivalves, gastropods, nautiloids and monoplacophorans with their originally aragonitic shells now dissolved. Many of the encrusting bryozoans are so thin and broad that they give the illusion of calcitic mollusc shells clinging to the moulds. Some molluscs in the Cincinnatian, especially monoplacophorans and epifaunal bivalves, would be poorly known if they had not been bryoimmured. Unlike internal and external moulds in sediment, bryoimmured fossils could be transported and thus record aragonitic faunas in taphonomic assemblages (e.g. storm beds) in which they would otherwise be rare or absent. In addition, bryoimmurations of aragonitic shells often reveal the ecological succession of encrustation on the shells by exposing the earliest encrusters and borings that were later overgrown. Bryoimmuration was common during the Late Ordovician because the calcite sea at the time quickly dissolved aragonitic shells on the seafloor before final burial, and large calcitic bryozoans very commonly used molluscs as substrates. Bryoimmuration is an important taphonomic process for preserving aragonitic faunas, and it reveals critical information about sclerobiont palaeoecology. Several Cincinnatian mollusc holotypes are bryoimmured specimens. Bryozoans involved in bryoimmuration enhance the preservation of aragonitic fauna and thus act as taphonomic engineers.     

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     

Bioerosional structures and pseudoborings from Late Jurassic and Late Cretaceous-Paleocene shallow-water carbonates (Northern Calcareous Alps, Austria and SE France) with special reference to cryptobiotic foraminifera

Examples of bioerosional processes (boring patterns) are described from shallow-water limestones of the Late Jurassic Plassen Carbonate Platform (PCP) and the Late Cretaceous to Paleocene Gosau Group of the Northern Calcareous Alps, Austria. Some micro-/macro-borings can be related to distinct ichnotaxa, others are classified in open nomenclature. In the Alpine Late Jurassic, bioerosional structures recorded from clasts in mass-flows allow palaeogeographical conclusions concerning the source areas. In particular, these are borings of the Trypanites-ichnofacies detected from clasts (Barmstein limestones) of the PCP or special type of bored ooids of unknown source areas or restricted autochthonous occurrences. In the Lower Gosau Subgroup, Gastrochaenolites macroborings occur in mobile carbonate clast substrates of shore zone deposits (“Untersberg Marmor”). Different types of borings are recorded from rudist shells and coral skeleton, some of which are referable to the ichnotaxon Entobia produced by endolithic sponges. In the present study, special attention is paid to the occurrences of the cryptobiotic foraminifera Troglotella incrustans Wernli and Fookes in the Late Jurassic and Tauchella endolithica Cherchi and Schroeder in the Late Cretaceous. The latter is so far only known to be from the Early Cenomanian of France and is reported here for the first time from the Late Turonian-Early Coniacian stratigraphic interval where it was found in turbulent carbonate deposits within borings penetrating bivalve shells or coralline algae. The records of cryptobiotic foraminifera from the Northern Calcareous Alps are supplemented by a single finding from the Middle Cenomanian of SE France. A palaeoenvironmental interpretation of the occurrences of the cryptobiotic foraminifera is provided.     

Depositional environment and constraining factors on the facies architecture of the Qom Formation,Central Basin,Iran

In the Central Iran Basin, the mixed carbonate–siliciclastic deposits of the C member of the Qom Formation were deposited on a carbonate platform which is dominated by rhodalgal associations occurring in tropical–subtropical environment. The biogenic rhodalgal association is dominated by bryozoa, coralline red algae, bivalves and echinoids together with smaller amounts of photo-dependent biota including large benthic foraminifera and corals. The abundance of heterozoan association and the bloom of suspension-feeding organisms are the result of an increase in nutrient availability which has profound controlling effect on the biotic system. The low occurrence of symbiont-bearing benthic foraminifera and coral, typical of stable, oligotrophic condition, represents their low tolerance to unstable, nutrient-rich environment. In the investigated Oligocene–Miocene shallow marine carbonate succession, 10 different microfacies were distinguished through depositional texture and biotic components. The rock sequences investigated are referred to an open shelf carbonate platform in which the depositional environments range from outer shelf to inner shelf conditions.     

Linking taphonomy to community-level abundance: Insights into compositional fidelity of the Upper Triassic shell concentrations (Eastern Alps)

Although actualistic live/dead comparisons lead to robust estimates of fidelity of modern death assemblages, quantitative evaluation of fidelity of fossil assemblage remains uncertain. In this paper, effects of storm reworking on compositional fidelity of the Upper Triassic shell concentrations (Eastern Alps, Austria) are evaluated. An exploratory approach is based on comparison of reworked and non-reworked assemblages in ordination analyses. Non-reworked assemblages of one or more communities provide a baseline for evaluation of fidelity of reworked assemblages. In siliciclastic-rich intervals of the Kössen Formation, shell concentrations are represented by (1) packstones with small, shallow infaunal bivalves, (2) floatstones and pavements with large semi-infaunal bivalves, and (3) bioclastic marlstones. In carbonate-rich intervals, bioclastic floatstones with bivalves and brachiopods occur. Analyzing all shell concentrations, eight sample groups sharing similar species composition are discriminated. Limited effect of storm reworking on composition of shell concentrations is indicated by (1) a general persistence of six sample groups when only non-reworked assemblages are analyzed, (2) similarity in composition between reworked and non-reworked assemblages within sample groups, and (3) compositional segregation between non-reworked assemblages of distinctive sample groups, mostly without any reworked assemblages of intermediate composition.Depth-related variations in dead-shell production, shell destruction and body size governed preservation and distribution of the shell concentrations along onshore-offshore gradient in the Kössen Basin. First, at times when environmental conditions were unfavorable for shell producers, coupled with high background shell destruction rates, limestone beds formed during storm events were shell-poor. Second, less common shell concentrations in upper than in lower parts of siliciclastic intervals can be related to higher environmental stress in shallower habitats. Third, the difference between shell concentrations dominated by small and large bivalves is driven by between-habitat differences in body size and is not due to a differential sorting of small and large shells. Combining community analysis based on species abundances with taphonomic analysis can thus be helpful in tracking fidelity of fossil assemblages.     

Encrustation patterns on Late Cretaceous (Turonian) echinoids from southern Poland

This study focuses on sclerobionts from a large collection of epibenthic echinoids (>2,000 specimens) of the genera Conulus and Camerogalerus. Samples were collected from five localities in southern Poland (Polish Jura and Miechów Trough), where Turonian carbonates with terrigenous input are exposed. Low intensity (mean ca. 5 %, maximum ca. 10 %) and slight encrustation (“loosening effect”) exclusively by episkeletozoans probably resulted from low productivity of encrusters while the importance of other factors cannot be excluded unambiguously. Echinoids served as a main substratum and after death formed shellgrounds (‘echinoid carpet’) offering abundant benthic islands for encrusters in an otherwise soft-bottom environment. The moderate abundance but low-diversity assemblage is represented by bivalves, sedentary polychaetes, foraminifera, bryozoans, corals, and sponges. This assemblage is similar to a nearly contemporaneous assemblage from the Bohemian Basin. The presence of numerous spirorbins offers insights into their early evolution and may indicate that their first peak in abundance after origination was not prior to the earliest Turonian. This is regarded as one of the important ecological steps towards the rise of modern sclerobiont communities. Encruster diversities are independent of their abundance and, as shown in our novel planar projections, lateral parts of tests were preferentially encrusted. This pattern is explained by the combination of largest flat area and stable orientation. Encrusting bivalves and serpulids dominated hard substrate environments in the Turonian of Poland.     

Strontium isotope stratigraphy of Cretaceous hippuritid rudist bivalves: rates of morphological change and heterochronic evolution

Strontium isotope stratigraphy of 17 localities of rudist formations in the region of the former Mediterranean Tethys has provided a reliable and precise stratigraphical frame for the evaluation of morphological change in hippuritid rudist bivalves during the Coniacian–Campanian. The phyletic lineage Vaccinites cornuvaccinum (Bronn)–Vaccinites chaperi (Douvillé) evolved from the Early Coniacian until the Early Campanian and is characterized by phyletic size increase and allometric growth, as shown by morphometrical measurements of 102 shells. These chronospecies intergrade in the Late Coniacian so that V. cornuvaccinum is considered to be a reliable marker species for the Coniacian. The taxonomy of Vaccinites alpinus (Douvillé) is discussed and the species is recognized as a senior synonym of Vaccinites ultimus (Milovanovi ). It appears first in the Late Santonian and the last appearance is probably in Late Campanian. Both lineages are characterized by phyletic size increase and peramorphic evolution involving hypermorphosis. A doubling of the length of the mantle margin occurred within 5 m.y. in both lineages. The results demonstrate that the combination of morphometric analyses and stratigraphical precision provides an important tool for the delineation of tempo and mode of evolution in rudist bivalves. Strontium isotope stratigraphy resulted in a considerable revision of the ranges of the species investigated. As the stratigraphy of many Tethyan carbonate platforms relies on the distribution of rudist bivalves, and the species investigated are abundant in many rudist formations, the history of many Late Cretaceous carbonate platforms must be re-evaluated.     

Bivalves with 'concrete overcoats': Granicorium and Samarangia

Two veneroidean bivalves Granicorium indutum from Australia and Samarangia quadrangularis from the tropical Indo-Pacific region, cement a thick, hard layer of sand over most of their shells. In Granicorium this layer forms low commarginal ribs while in Samarangia it forms more prominent radial features. Sand grains are cemented to the shell and to each other with growths of a crystalline aragonitic cement similar in morphology to inorganic marine cements. Both species secrete mucus layers at the growing shell margin which initially hold the sediment grains together and form a substrate for the nucleation and growth of calcium carbonate crystals. The ribs of Samarangia are formed by the accretion of successive sheets of spherulitic growths. In G. indutum , the middle and outermost of two inner mantle folds are large, glandular and capable of considerable extension beyond the shell margin. Mucus secreted by the folds contains abundant bacteria and small calcium carbonate crystals. It is proposed that initial nucleation of the calcium carbonate cement takes place within this biofilm possibly mediated by the bacteria. The function of the sand layers is unknown but predation resistance and protection of the shells from endobionts are the most likely possibilities.     

Mixed siliciclastic–cool-water carbonate deposits over a tide-dominated epeiric platform: the Faluns of l’Anjou formation (Miocene,W. France)

The Tortonian bioclastic sands of Anjou (W France) are weakly cemented mixed siliciclastic–carbonate deposits. The carbonate fraction can reach up to 90%, and is irregularly spread over the area of the original depositional platform. The temperate marine water character is demonstrated by the lack of ooids, green algae, and biohermal scleractinians, and is dominated by numerous species of bryozoans and bivalves, associated with red algae, barnacles, and echinoderms (bryomol facies; Heterozoan association). Skeletal grains are weakly cemented. The presence of large submarine dunes indicates a platform bounded by a tide-domination of a succession that developed rapidly under highstand and shelf margin wedge system tracts. The tide-dominated “Faluns de l’Anjou” provides a model different from many other examples of temperate carbonate settings, which are often wave-dominated.     

Facies analysis of a large-scale Jurassic shelf-lagoon: the Kamar-e-Mehdi Formation of east-central Iran

The Callovian–Lower Kimmeridgian Kamar-e-Mehdi Formation of the Tabas Block (east-central Iran) is an up to 1,350-m-thick, fine-grained, marly-calcareous unit containing a basal Echellon Limestone Member (up to 180 m thick) and a terminal Nar Limestone Member (up to 100 m thick). The formation was deposited in a relatively deep shelf-lagoon that was part of the large-scale carbonate system of the Esfandiar Subgroup, extending N–S for about 500 km along the strike with a width of up to 100 km. The lagoonal Kamar-e-Mehdi Formation shows sedimentation rates of 150 m/myr, twice as high as those of the shelf-edge carbonate barrier (Esfandiar Platform). The repetitive lithologies and uniform depositional environment suggest equilibrium conditions between sedimentation and subsidence, related to constant slow rotation of the Tabas fault-block around a horizontal axis, the platform sitting on the crest, and the lagoon occupying the dip-slope. Lagoonal sedimentation was dominated by suspended carbonate mud and peloids from the eastern Esfandiar Platform whereas the subordinate siliciclastic material was derived from the west (Yazd Block). The diverse macrobenthos (mainly bivalves) suggests fully marine conditions for the major part of the Kamar-e-Mehdi Formation. However, towards the upper part, biotic impoverishment and the deposition of skeletal-poor, evaporitic sediments indicate increasing restriction. The overlying Magu Gypsum Formation marks the end of an arid basin-fill cycle and possibly forms an effective seal for hydrocarbon reservoirs in that area. The Esfandiar Subgroup was a Neotethys-facing carbonate margin, forming part of a belt of carbonate systems tracking the margins of the Iran Plate during Callovian to Late Jurassic times.     

北羌塘盆地中南部侏罗纪布曲组古生物特征与沉积环境

本文通过对北羌塘盆地北坳陷中南部胜利河、东湖及毛毛山等地区侏罗纪布曲组6条剖面及其中丰富的腕足类、双壳类等古生物资料和岩石组合特征的研究,将布曲组地质时代划为中侏罗世巴通期(Bathonian)至早卡洛夫期(Callovian),还可能跨入早巴柔期(Bajocian)。通过本文研究和区域对比,认为布曲组的沉积时代在北羌塘盆地存在穿时性。依据岩石组合特征,布曲组沉积充填物三分性明显,下部和上部为一套以微晶结构、粒泥结构为主的低能碳酸盐岩,中部为一套以高能的亮晶粒屑灰岩为主的碳酸盐岩。结合古生物生态习性,布曲组沉积环境总体为近岸浅水开阔台地–台地边缘碳酸盐岩沉积体系,构成多个沉积旋回。这一基础资料对下一步分析北羌塘盆地坳陷中南部布曲组岩相古地理提供了支撑。     

Paleoecology of epizoans and borings on some Upper Cretaceous chalk oysters from the Gulf Coast

Analysis was made of a hard substrate fauna found on right valve interiors and exteriors of the epifaunal reclining oyster Pycnodonte mutabilis from the Maastrichtian (Navarroan) Saratoga Formation (southwestern Arkansas). Comparison of boring and encrustation patterns on both sides of valves indicates that a major portion of colonization on valve exteriors occurred while host oysters were alive. Paleoautecologic information derived from such valve exterior patterns includes evidence of rheotropic orientation by encrusting juvenile P. mutabilis and preferential location of Trypanites sp. borings in surficial shell grooves. Valve exteriors supported a hard substrate paleocommunity which had the following non-interactive progressive colonization sequence: (1) Trypanites sp. and P. mutabilis juveniles; (2) Entobia sp., serpulid worm tubes, and Bullopora sp.; and (3) cheilostome bryozoans. This sequence could have been caused by low seasonality and ranked success of colonizing encrusters and borers. Colonization of valve interiors generally differed from exteriors only in that many interiors were first colonized by the clionid sponge that created Entobia sp., which had already occupied the exterior, and which quickly bored through the valve to occupy the interior upon the host's death. □ Trace fossils, epizoans, borings, Gryphaeidae, palaeoecology, communities, colonization sequence, Late Cretaceous, Maastrichtian, Navarroan, Arkansas.