Determination of a Late Miocene rocky palaeoshore by bioerosion trace fossils from the Bozcaada Island, Çanakkale,Turkey

Bioerosion is a common process in hard substrates. This study introduces an example from the rocky palaeoshore cropping out at a sea cliff on the Bozcaada Island. It includes bioerosion trace fossils preserved in limestone boulders of the shallow marine and lacustrine Alcitepe Formation of Late Miocene age. The ichnotaxa include borings produced by duraphagous drillers (Oichnus isp.), phonorids (cf. Conchotrema isp.), clionid sponges (Entobia cf. goniodes, Entobia geometrica, Entobia laquea, Entobia ovula, E. cf. solaris, Entobia isp.), endolithic bivalves (Gastrochaenolites torpedo, Gastrochaenolites lapidicus, Gastrochaenolites isp., Phrixichnus isp.), polychaete annelids (Maeandropolydora isp., Maeandropolydora sulcans, Maeandropolydora decipiens, Caulostrepsis taeniola, Caulostrepsis isp.), echinoids (cf. Circolites isp.) and spinculid worms (cf. Trypanites isp.). Barnacles are also common as encrusters. The borings can be ascribed to the Gastrochaenolites-Entobia assemblage, which is typical of Neogene rocky-shores. They belong to the Entobia ichnofacies indicating various conditions of light, energy, and depth. Therefore they can reveal environmental changes and play an important role in forming palaeo-rocky shores and wave-cut platforms during marine trangressive events.     

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).     

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.     

Devonian Trace Fossils of the Horlick Formation,Ohio Range,Antarctica: Systematic Description and Palaeoenvironmental Interpretation

The clastic Horlick Formation contains an ichnofauna of 28 ichnotaxa dominated by burrowers. These are a mixture of simple vertical forms (Skolithos linearis, S. magnus, Bergaueria cf. langi, Rosselia socialis, Monocraterion isp.), U or arc-like forms (Diplocraterion parallelum, Arenicolites types A and B, Catenarichnus antarcticus, C. isp., aff. Lanicoidichna isp.), and complex, vertical spiral structures (Asterosoma isp., Spirophyton isp.). Horizontal burrows include Ancorichnus cf. capronus, Palaeophycus tubularis, and Psammichnites devonicus isp. nov. Surface traces comprise Haplotichnus isp., Cruziana problematica, C. rhenana, Rusophycus aff. carbonarius, R. isp., Protovirgularia rugosa, Lockeia ornata and cubichnia indet., while trackways include Diplichnites gouldi, D. isp., Maculichna? isp. and large imprints. The Horlick Formation (maximum 56 m) records an early Devonian transgression onto a deeply weathered land area that lay in the direction of Marie Byrd Land, spreading from the South Africa sector of Gondwana. Analysis of the trace fossils confirms their formation in near-shore to intertidal environments, with some ichnotaxa living close to the marine/fluvial boundary (e.g., Spirophyton, aff. Lanicoidichna, Cruziana problematica, Rusophycus aff. carbonarius, Maculichna?). The ichnofauna probably comprised a variety of marine suspension and deposit-feeding worms, shallow burrowing molluscs (bivalves and possibly bellerophontids) and several different types of arthropods, including trilobites.     

Rocky-shore unconformities marking the base of Badenian (Middle Miocene) transgressions on Mt. Medvednica basement (North Croatian Basin,Central Paratethys)

Badenian (Middle Miocene) transgressive deepening-upward successions located in the NE part of Mt. Medvednica (North Croatian Basin, Pannonian Basin System) unconformably overlie Mesozoic basement. Triassic and Upper Cretaceous limestone pebbles, cobbles, and boulders of the Badenian basal conglomerates display abundant in situ bivalve borings of Gastrochaenolites and sponge borings of Entobia. This Gastrochaenolites-Entobia ichnoassemblage is related to the Entobia subichnofacies of the Trypanites ichnofacies, characterizing littoral rocky-shore environments (wave-cut platforms and marine transgressive surfaces with a low or null rate of sedimentation). Gastrochaenolites torpedo, Gastrochaenolites lapidicus, and Entobia recorded in Badenian basal conglomerates (compared with modern Northern Adriatic rocky-shore environments), enabled more precise palaeoenvironmental interpretations. The occurrence of G. torpedo (produced by lithophaginid bivalves) on all sides of individual limestone lithoclasts in the Gornje Ore?je basal conglomerate, coupled with truncation of the formation (possibly indicating multiphase colonization), reflect gravel transport, roll-over, overturning and erosion by wave action in high-energy rocky-shore settings. Gornje Psarjevo-2 basal conglomerate boulders were probably not subjected to significant movement and abrasion, as suggested by good preservation of both G. lapidicus (potentially produced by gastrochaenid bivalves), associated G. torpedo, as well as abundant shallow Entobia borings. The Badenian Gastrochaenolites-Entobia ichnoassemblage also could be the result of a composite development. However, direct cross-cutting relationships between G. torpedo and G. lapidicus and/or Entobia were rarely observed. In addition, Badenian boring tracemakers might have coexisted at the same water depth. Northeast Mt. Medvednica Badenian successions probably formed during different Central Paratethys transgressive pulses (NN5 and NN6 Zones). However, exact timing of Badenian transgressions, stratigraphic correlations and tectono-eustatic implications are unresolved, due to sparsely integrated biostratigraphic and high-precision geochronological studies of Early–Middle Miocene North Croatian Basin deposits and due to the absence of a uniform biostratigraphic zonation and regional chronostratigraphic division of Central Paratethys.     

A Distinctive Bioglyph and its Producer: Recent Gastrochaenolites Leymerie in a Peat Pebble,North Sea Coast of The Netherlands

A pebble of Quaternary peat collected from the Dutch coast north-northeast of Zandvoort, Noord Holland, preserves a recent boring, Gastrochaenolites isp. cf. G. lapidicus. This is geniculated and retains a distinctive, corrugated bioglyph. The probable producing bivalve was the pholad Barnea candida (Linné), known as the white piddock, or Petricola pholadiformis Lamarck, known as the American piddock, identified by beachcombing for boring shells. But these form a minute numerical fraction of a modern shelly death assemblage dominated by burrowing bivalves; an analogous approach with the fossil record is less likely to succeed.     

Ichnofacies and microbial build-ups on Late Miocene rocky shores from Menorca (Balearic Islands), Spain

Angular unconformites between Jurassic and Miocene strata are exposed in the sea cliffs of Cala Cigonya on the northwest coast of Menorca in the Balearic Islands of Spain. The geological discontinuities represent rocky shores on opposite sides of a former headland with 15 m of topographic relief. On the east flank, Jurassic dolomite is overlain by Upper Miocene (Tortonian to Messinian) breccia and laminated limestone. Here, a partially exhumed dolomite surface records Miocene bivalve borings of the ichnospecies Gastrochaenolites torpedo and G. lapidicus that achieved a density of >1,000 borings/m². Other associated traces include sponge borings (Entobia isp.) and polychaete borings (Caulostrepsis isp.). A breccia deposit 0.8 m thick was derived from the underlying dolomite and angular clasts still retain evidence of bivalve borings. Above follows a succession of laminated limestone beds more than 5 m thick, including some levels with dome-shaped stromatolites and other horizons with reworked dolomite clasts. Thin-section analysis of the laminated limestone reveals dark and light couplets 0.2 mm thick consistent with microbial origins. In contrast, the west flank was buried by coarse sandstone and laminated sediments. Here, dwelling structures of regular echinoids (Circolites kotoncensis) are the dominant traces preserved on the dolomite surface, reaching a maximum density of 66 borings/m². Associated borings include Entobia geometrica as well as rare traces of Gastrochaenolites isp. and Trypanites isp. Notable for the absence of a basal Miocene breccia, the west flank is interpreted as a sheltered rocky shore coeval with an exposed rocky shore on the east flank. Today, heavy surf on the north coast of Menorca is related to the Tramontana winds that blow out of Spain during winter months. Similar atmospheric circulation patterns must have prevailed during the Late Miocene, but the replacement of ichnofacies by microbial build-ups resulted from increased salinity during the Messinian crisis.     

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.     

Evidence of Bioerosive Structures in Quaternary Glaciomarine Sediments from Southwestern Iceland

The Late Weichselian marine and glaciomarine sediments occurring in many places along the coast of Iceland are rich in invertebrate fossils, especially mollusks and barnacles. A diverse ichnofauna, constituted especially by bioerosional traces documenting the activity of predators and animals that use the host shell for attachment or the construction of dwellings, was identified at the localities of Ósmelur, Saurbær, and Brekkubakkar in southwestern Iceland. The ichnotaxa identified include Anellusichnus circularis (Santos, Mayoral, and Muñiz), Caulostrepsis isp., Centrichnus concentricus (Bromley and Martinell), Clionolithes isp., Finichnus peristroma (Taylor, Wilson, and Bromley), Finichnus isp., Sedilichnus asperus (Nielsen and Nielsen), Sedilichnus cf. excavatus (Donovan and Jagt), Sedilichnus gradatus (Nielsen and Nielsen), Sedilichnus smiley isp. nov., Sedilichnus ovalis (Bromley), Sedilichnus paraboloides (Bromley), Sedilichnus simplex (Bromley), and Sedilichnus spongiophilus (Müller).     

The infaunal echinoid Micraster: Taphonomic pathways indicated by sclerozoan trace and body fossils from the Upper Cretaceous of northern Spain

Micraster echinoid fossils are common in the Upper Cretaceous Olazagutía Formation of northern Spain. Tests frequently record sclerobiont signatures (including bioerosion and encrustation), left by reaction and/or coaction phenomena. Among bioerosion structures, Oichnus simplex, O. paraboloides, O. ichnosp. A, Trypanites solitarius, Rogerella ichnosp. indet., Centrichnus cf. eccentricus, Maeandropolydora ichnosp. indet. and fungal microborings are found, as well as pits and fractures. As for sclerozoan body fossils, bivalves (Dimyidae, Anomiidae, Plicatulidae and other Pectinacea), polychaete annelids (serpulids and spirorbids), lituolid foraminiferans (Haddoniidae and Coscinophragmatidae) and bryozoans (cheilostomate ones and others undetermined), as well as other less common groups, have been identified. Taphonomic paths followed by Micraster tests are analysed, based on conservation state and degree of colonisation and sedimentary filling. This allows to distinguish between accumulated fossils and non accumulated ones (including resedimented and reelaborated ones).     

Composite Trace Fossil Assemblage in a Distal Carbonate Setting from the Tethys (Middle Jurassic,Betic Cordillera,Southern Spain)

Ichnological analysis of a Middle Jurassic carbonate surface from the Betic Cordillera (southern Spain) reveals a complex trace fossil assemblage, including softground Ophiomorpha, firmground Arenicolites, Thalassinoides and Gastrochaenolites, and hardground Trypanites as well as possible Gastrochaenolites. The ecological replacement in the macrobenthic community is interpreted according to successive suites that are controlled mainly by substrate consistency. Variations in composition and abundance of trace fossils between suites can be ecologically and/or taphonomically determined.     

Bored and Burrowed: An Unusual Echinoid Steinkern from the Type Maastrichtian (Upper Cretaceous,Belgium)

A test of the irregular echinoid Hemipneustes striatoradiatus (Leske) was dissolved in dilute hydrochloric acid to expose the flint steinkern. This revealed slender, string-like borings that were mainly limited to ambulacra (Talpina isp. cf. T. canna (Price)); less common pouch-like borings limited to part of one ambulacrum only (Rogerella isp. cf. R. pattei (Saint Seine)); and slender, sinuous to contorted burrows, Planolites montanus Richter. Ingress to the test (borers) and sediment infill (burrowers) was presumably through the ambulacral pore pairs of the dead test. Borings and burrows were probably not coeval. Such infestations of fossil echinoids may be locally common, but their identification is controlled by the difficulty of seeing features of both the test and the infill.     

A New Ichnospecies of Gastrochaenolites Leymerie from the Pleistocene Port Morant Formation of Southeast Jamaica and the Taphonomy of Calcareous Linings in Clavate Borings

The ichnospecies Gastrochaenolites pickerilli isp. nov. is based on ten borings found in a shell of the gastropod Strombus gigas Linné from the Pleistocene (Sangamonian) Port Morant Formation of southeast Jamaica. These borings bear morphological similarities to Gastrochaenolites torpedo Kelly and Bromley but differ from all other Gastrochaenolites ispp. in having prominent and numerous calcareous meniscate structures arrayed adjacent to one side of the boring. These menisci are concave towards the center of the boring and are the remnants of calcareous tubes that lined earlier boreholes, that the boring bivalve treated as part of the lithified substrate when relocating. They are thus evidence of the former positions of borings that, unusually, were breached as the bivalve migrated sideways. Although this was a common behavior for Gastrochaenolites-producing bivalves within this substrate, the reason for it occurring is uncertain.     

Bioerosion in the Miocene Reefs of the northwest Red Sea,Egypt

Macroborings provide detailed information on the bioerosion, accretion and palaeoenvironment of both modern and fossil reefs. Dolomitized reefal carbonates in the Um Mahara Formation exhibit an outstanding example of spatially distributed, well‐preserved bioerosion structures in tropical to subtropical syn‐rift Miocene reefs. Ten ichnospecies belonging to five ichnogenera are identified; three belonging to the bivalve‐boring ichnogenus Gastrochaenolites, three attributed to the sponge‐boring ichnogenus Entobia, and four ichnospecies assigned to three worm‐boring ichnogenera Trypanites, Maeandropolydora and Caulostrepsis. The distribution of the reported borings is strongly linked to the palaeo‐reef zones. Two distinctive ichnological boring assemblages are recognized. The Gastrochaenolites‐dominated assemblage reflects shallower‐marine conditions, under water depths of a few metres, mostly in back‐reef to patch‐reef zones of a back‐reef lagoon. The Entobia‐dominated assemblage signifies relatively deeper marine conditions, mostly in reef core of the fringing Miocene reefs. These ichnological assemblages are attributed herein to the Entobia sub‐ichnofacies of the Trypanites ichnofacies. This ichnofacies indicates boring in hard carbonate substrates (such as corals, rhodoliths, carbonate cements and hardgrounds) during periods of non‐sedimentation or reduced sediment input.     

Holes in Bones: Ichnotaxonomy of Bone Borings

Bone is a substrate for bioerosion at equal rank with xylic and lithic substrates. Accordingly, borings in bone have to be identified in an analogous way to other ichnogenera coined for one type of substrate. In due course, the new ichnogenera Osteichnus n. igen. and Clavichnus n. igen. are established within the new ichnofamily Osteichnidae. Gastrochaenolites and Trypanites are here restricted to lithic substrates, and Asthenopodichnium only occurs in xylic substrates. Only with this approach, ichnotaxobases of trace fossils in bone are identical to those in other hard substrates. Cuniculichnus variabilis n. igen. n. isp. is introduced for variably shaped pits to tunnels bored into bone by beetle (arguably dermestid) larvae; its ethological character is close to a pupichnion.     

New bivalve burrows from the mid-Holocene of northeastern Buenos Aires Province (Argentina): ichnotaxonomy and ethology

Oblongichnus vulpesi n. isp. is herein described as amygdaloid, oval or cleft-shaped burrows with a lining of variable thickness and an oblong to quadrangular elongated shaft from the mid-Holocene of the Destacamento Río Salado Member. Valves of the solenid bivalve species Solen tehuelchus were found within the burrows, indicating that this razor clam is the tracemaker. This new finding corroborates the validity of Oblongichnus and previous interpretation that the burrow was produced by infaunal, filter feeding bivalves with a foot of very limited horizontal movements. We found that the origin of oblong elongate bivalves may be dated back to the Ordovician, while an infaunal radiation likely occurred during late Paleozoic. We interpret that O. vulpesi was made in a moderate energy subtidal marine setting during the last marine highstand.     

Epi- and endobiontic organisms on Late Jurassic crinoid columns from the Negev Desert, Israel: Implications for co-evolution

Columns of the articulate crinoids Millericrinus and Apiocrinites from the Upper Jurassic (Upper Callovian) Zohar and Matmor formations of the Negev Desert of Israel display abundant encrusting organisms of about ten species, as well as diverse trace fossils produced by endobionts. Pluricolumnals were colonized by epi- and endobiontic organisms both during life and post-mortem. Skeletonized encrusting organisms include abundant ostreid bivalves (which evidently colonized both live and dead crinoid columnals), two types of serpulid worms, encrusting foraminifera, three species of bryozoans, and small encrusting sclerosponges. Several types of borings are present: Trypanites (possibly produced by sipunculids), Gastrochaenolites (crypts of boring lithophagid bivalves), elliptical barnacle? borings, and channel-like annelid? borings. In addition, approximately 16% of the pluricolumnals display circular parabolic embedment pits assignable to the ichnogenus Tremichnus. They are associated with substantial deformation of the containing columnals and were probably the work of host-specific ectoparasitic organisms. Discovery of Tremichnus on Jurassic crinoids extends the range of this trace by almost 100 million years, providing evidence for one of the longest-ranging host-parasite interactions documented thus far (over 200 million years). The relationship of epibionts to the Jurassic crinoids thus ranged from simple utilization of dead hard substrate to probable opportunistic commensalism in forms that colonized the live upright stems, as in some oysters, through host-specific parasitism in the case of Tremichnus.     

Effect of depth and reef structure on early macrobenthic communities of the Algarve artificial reefs (southern Portugal)

This study was carried out on the “Faro/Ancão” artificial reef (AR), located off Faro, deployed in May 2003. We aimed to characterise early macrobenthic community colonisation of two concrete AR groups located at different depths (16 m and 20 m depth) and to test the effect of reef structure on these communities. The non-colonial organisms were counted; barnacles and colonial species were quantified using biomass. Multivariate analyses indicated that early macrobenthic communities (6 months of immersion) were affected by depth, and that barnacles and colonial species were also affected by reef structure. Univariate analyses showed that the biomass of barnacles and colonial species was significantly different among reefs and layers of modules. Both AR groups were characterised by the species Balanus amphitrite, Gregariella subclavata, Musculus cf. subpictus, Paleanotus cf. bellis and Syllidia armata. Jassa marmorata and Bugula neritina were characteristic species at 16 m depth, particularly on the AR Upper layer of modules, whereas Anomia ephippium was particularly common at 20 m, especially on the Lower layer of modules.     

A new possible bivalve burrow Oblongichnus solodukhoi from the late Kazanian (middle Permian) stratotype section in Russia

A burrow previously assigned to the ichnotaxon Palaeophycus insignis is redescribed, and a new ichnotaxon Olongichnus solodukhoi n. igen. n. isp is erected for this flattened, slightly curved, multi-oriented, smooth burrow that has a thick lining with internal and external very fine-grained mucuous layers. It shows subrectangular to oblong form in cross section. O. solodukhoi is collected from the lowermost part of the Member D (Seryi Kamen Member, Pechishchi Beds) of the late Kazanian (middle Permian) stratotype section (East European Platform). It occurs in yellowish grey dolostone formed within open shallow-marine environments. O. solodukhoi records combined feeding and dwelling behaviours of a suspension-feeding organism. It could be produced by burrowing organisms with a simple flattened morphology, such as flatworms or polychaetes; however, we favour elongate and ultra-elongate bivalves as the potential tracemakers of the reported burrows.     

Investigation of toxin profile of Mediterranean and Atlantic strains of Ostreopsis cf. siamensis (Dinophyceae) by liquid chromatography–high resolution mass spectrometry

Blooms of Ostreopsis spp. once confined to tropical and subtropical areas have recently spread to more temperate regions such as the Mediterranean and the Southern-Atlantic coasts of Europe. However, while O. confronta (cf.) ovata has caused several toxic outbreaks, the presence of O. cf. siamensis has been reported rather occasionally and in very few regions; as a consequence, O. cf. ovata toxin profile has been in-depth studied while poor information exists on toxicity of the Mediterranean and Atlantic O. cf. siamensis. In the present study toxin profile of Mediterranean and Atlantic O. cf. siamensis isolates also phylogenetically related has been studied through liquid chromatography–high resolution mass spectrometry (LC–HRMS) versus a palytoxin standard, a crude extract of O. cf. ovata containing all the ovatoxins so far known (ovatoxin-a to -f), and a Japanese O. siamensis extract which contained ostreocin-d and ostreocin-b. The Mediterranean and Atlantic O. cf. siamensis strains were shown not to produce either ostreocins, which are produced by the Japanese O. siamensis strain, or ovatoxins, which are produced by the Mediterranean O. cf. ovata. Only sub-fg levels of palytoxin on a per cell basis were detected in the Mediterranean strain. This study demonstrates that the Mediterranean and the Atlantic O. cf. siamensis strains are devoid of any appreciable toxicity. Thus, at least in the European area, O. cf. siamensis seems to present a much lower risk to human health than O. cf. ovata.