Boring Late Cambrian organisms

Specimens of inarticulate brachiopods (family Acrotrctidae) with boreholes were found in Upper Cambrian carbonates in the southern Great Basin of the United States. Some morphologic features and preferred orientation of the borings are similar to those made by predators and previously reported in the fossil and Holocene record. Such predatory activity on brachiopods is previously unknown in Cambrian rocks. Taxa associated with these specimens are not known to have been predators and identity of the predatory organism is unknown. Cambrian brachiopods, predation, paleoecology.     

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.     

Predatory borings in the inarticulate brachiopod Artiotreta from the Silurian of Oklahoma

A sample containing inarticulate acrotretid brachiopods, from the Silurian Clarita Formation of Oklahoma, was studied to determine the occurrence, nature and frequency of boring in the shells, and the possible identity of the boring organism. Specimens of Artiotreta parva Ireland, 1961 are abundant and specimens of Opsiconidion n. sp. and Acrotretella siluriana Ireland, 1961 are rare in the samples studied. All of the borings discovered are in A. parva except for one, which is in a specimen of O. n. sp. The boreholes are considered to be of predatory origin, and are similar in morphology to borings made by Recent gastropods. The small countersunk boreholes are equally frequent on pedicle and brachial valves, but preferentially located in particular regions of each valve. Boring frequency is high. The identity of the boring organism has not been determined, but is considered to be a soft bodied predator or an archaeo-gastropod.     

Recent borings in limestone cobbles from Marloes Bay, southwest Wales

Limestone clasts from the beach at Marloes Sands, southwest Wales, contain slender, straight to sinuous borings cross-cut by younger, clavate borings. The former were probably produced by sipunculids or polychaetes; the latter preserve shells of the boring bivalve Gastrochaena dubia (Pennant). Unusually, the calcareous linings of the clavate bivalve borings extend into many of the slender worm borings. Such linings are considered part of the hard parts of the producing bivalve, but the chance association of the two morphologies of borings has led to the lining becoming intimately associated with both of them. The modified linings of the bivalve borings have a similar morphology to the crypt of certain clavagellid bivalves, perhaps presenting an analogue for the morphology of a pre-clavagellid, boring ancestor.     

Microboring in a Freshwater Fluvial Unionid Bivalve Substrate

Samples of the unionid bivalve Elliptio complanata were collected from the channel of the freshwater Saint John River, from Fredericton, New Brunswick, Canada. Scanning electron microscopy imaging of prepared shell samples revealed an assemblage of microborings. No borings are noted on the periostracum or prismatic shell layers. Boring structures are instead confined to the underlying nacreous aragonitic shell material, together with its associated organic conchiolin layers. Three main styles of boring are encountered, encompassing both predominantly surficial structures and penetrative tubular borings. Surficial structures are represented by a polygonal network on an exposed conchiolin shell layer. The penetrative borings take two forms, one being simple unbranched tubes, steeply aligned (perpendicular to the shell surface) and traversing the full thickness of the nacreous shell layer. The other penetrative boring style, again occurring within the nacreous layer, comprises a complex irregular network of randomly oriented rarely branching tubular borings. Borings generally display diameters of micron scale. Biofilm and extracellular polymeric substances, with bacterial, diatomaceous and filamentous components are also observed, often displaying a close association with both the borings and the conchiolin layers within the shell. The formation of the borings may be attributed to cyanobacteria, cyanophyte or fungal progenitors.     

Parasitism on Ordovician Chitinozoa

Borings are common in chitinozoans. They are cylindrical or conical, widening towards the outer surface of the vesicle wall. It is suggested that the cylindrical borings were probably caused by bacteria. Actinomycota and lower Fungi including at least Chytridiomycota, and the conical borings mainly by chytridia. Two types of spore-like bodies, operculate and non-operculate, have been found in situ on two chitinozoan species. The spore-like bodies are of about the same size as the supposed boring organisms in chitinozoans but are never found in association with the borings. They are interpreted as cysts of ectoparasitic nannoorganisms that resemble ciliates and amoebae, and their possible use for determining the biochemical composition of the chitinozoans is briefly discussed.     

Evidence of Osedax worm borings in Pliocene (∼3 Ma) whale bone from the Mediterranean

Osedax worms subsist entirely on vertebrate skeletons on the seafloor, using root-like tissues to bore into and degrade the bones. Paleontologists have only recently begun to appreciate the possible destructive effect that these worms may have had on the marine vertebrate fossil record and little is known of their evolutionary history. Using microcomputed tomography, we document Osedax-like borings in a fossil whale bone from the Pliocene of Italy and present new data on the borings of extant Osedax worms. The fossil borings are distinguished from those of other known borers and identified as traces of Osedax activity based on diagnostic features. Our results suggest that it is necessary to isolate individual borings for the confident identification of Osedax traces. This is only the second paleogeographic occurrence of Osedax in the fossil record and indicates that by the Pliocene these worms had colonised a large portion of the world's oceans. This is the first evidence for Osedax in the Mediterranean, past or present, and suggests that more species await discovery in this region.     

Bivalve Wood Borings of the Ichnogenus Teredolites Leymerie from the Bohemian Cretaceous Basin (Upper Cretaceous,Czech Republic)

Finds of fossil wood with bivalve wood borings (Teredolites clavatus and T. longissimus) occur in various facies and presumed sedimentary settings of the platform, shallow-marine Bohemian Crectaceous Basin. The basin comprises areas with sandy-dominated sediments, with marl and clay-dominated sediments, areas with predominat sandy-marly rocks, and finally areas dominated by calcareous nearshore sediments. Teredolites clavatus is common in fossil wood of sandstones, originating in beach or deltaic settings; marl and clay-dominated rock frequently bear wood fragments densely bored by Teredolites longissimus. When accompanied by evidence of marine environments as body fossils, glauconite or typical trace fossils, most of the wood fragments are bored. The presence/absence of borings in wood fragments can be considered the most reliable and easily useable criterion of distinction of marine settings in sandy sediments of the margin of the Bohemian Cretaceous Basin.     

Male golden hamsters (<Emphasis Type="Italic">Mesocricetus auratus</Emphasis>) are more reactive than females to a visual predator cue

In most species of small mammals, males are exposed to higher levels of risk than females because they compete for mates, travel greater distances to find and procure mates, and/or defend a territory. This suggests that males and females might have different responses to risky situations, such as the presence of a predator. We tested responses to a visual predator cue (an owl silhouette) in male and female golden hamsters (Mesocricetus auratus). In a laboratory arena, there was no significant sex difference in the latency to enter the burrow or time spent in the burrow immediately after exposure to the owl silhouette. Males, however, were less likely to be active during the 3-min period following the animal’s exposure to the silhouette, indicating that male golden hamsters are more wary after exposure to an aerial predator cue than females. Most studies of responses to predators or predator cues have not considered sex differences, but our results show that males and females may have quite different responses to predator cues. Further work should be done to characterize and quantify sex differences in response to predators or predator cues.     

Origin and paleoecology of Middle Jurassic hiatus concretions from Poland

Bored and encrusted carbonate concretions, termed hiatus concretions, coming from the Middle Jurassic (Upper Bajocian and Bathonian) siliciclastics of the Polish Jura, south-central Poland, have been subjected to detailed paleoecological investigation for the first time. The concretions possess variable morphology and bear distinct traces of bioerosion and encrustation as a result of exhumation on the sea floor during intervals of low sedimentation and/or erosion. The borings are dominated by Gastrochaenolites and Entobia. Epilithozoans, represented by at least 26 taxa, are dominated by sabellid/serpulid worm tubes and bryozoans, while sponges and corals are minor. No relationship between the concretion size and the number of encrusters has been found, suggesting that concretion size was not the primary factor controlling diversity. Stable isotope analyses and the presence of crustacean scratch marks and Rhizocorallium traces on many of the hiatus concretions indicate that they formed just below the sediment–water interface, within the sulfate reduction zone. Moreover, crustacean activities may have been a prelude to their origin, as shapes of many concretions closely resemble thalassinoidean burrow systems. It is also possible that crustacean activity around the concretions promoted their exhumation by loosening the surrounding soft sediment. The presence of borings and encrusters on different concretion surfaces, as well as truncated borings and a number of abraded epilithozoans, indicate that after the concretions were exhumed they were repeatedly overturned and moved on the sea floor, probably due to episodic storm-related bottom currents in shallow subtidal environment.     

Host‐specific acrothoracid barnacles on Middle Devonian platyceratid gastropods

Borings, attributed to acrothoracic barnacles, occur on the platyceratid gastropod Naticonema lineatum (Conrad) from the Middle Devonian Hamilton Group of western New York and rarely in specimens as old as the Early Devonian. These latter are the oldest known acrothoracid borings are in the fossil record. The borings are consistently developed as laterally compressed, inequilateral pouches exclusively on these gastropods, commonly as dense infestations.

Naticonema shells yielding borings typically occur associated with partially articulated qrinoid remains, and they are sometimes found attached to crinoids in a manner similar to coprophagous Platyceras. In addition to barnacle borings, Naticonema shells often bear thin encrustations of bryozoans which are usually perforated by these borings but sometimes also overgrow them. Barnacles bored live hosts; gastropods prevented shell penetration by producing cyst‐like secondary secretions of calcite beneath acrothoracid boreholes.

The relative antiquity of these borings and their association with coprophagous platyceratids makes their discovery particularly significant in revealing aspects of the early ecology of barnacles. Attachment to the host commensal gastropods was one of the first successful life modes of these crustaceans prior to their later diversification to other habitats. Mississippian and Pennsylvanian occurrences of similarly bored gastropods demonstrate continuity of the barnacle‐gastropod‐crinoid ecological association from the Middle to Late Paleozoic.     

Site selection and behavior of sponge and bivalve borers in shells of the cretaceous oysters exogyra cancellata and pycnodonte mutabilis from delaware,U.S.A

In shells of the oysters Exogyra cancellata and Pycnodonte mutabilis from the Mount Laurel and Marshalltown Formations (Campanian‐Maestrichtian), three‐quarters of all valves bear sponge borings (Entobia isp.) borings and 30% have borings of a lithophagid bivalve (Gastrochaenolites isp.). Non‐random distributions of these euendoliths, documented in this paper, may in part be accounted for by differential survival of sponge and lithophagid larvae and spat in varying circumstances. In addition, exterior shell architectures and post‐mortem orientations of shells are inferred to have prompted active geophobic (antigravity), rugophilic (groove‐seeking), and rheophilic (current‐seeking) behavior that enhanced survivorship of the settling larvae.     

Dynamics of a intraguild predation model with generalist or specialist predator

Intraguild predation (IGP) is a combination of competition and predation which is the most basic system in food webs that contains three species where two species that are involved in a predator/prey relationship are also competing for a shared resource or prey. We formulate two intraguild predation (IGP: resource, IG prey and IG predator) models: one has generalist predator while the other one has specialist predator. Both models have Holling-Type I functional response between resource-IG prey and resource-IG predator; Holling-Type III functional response between IG prey and IG predator. We provide sufficient conditions of the persistence and extinction of all possible scenarios for these two models, which give us a complete picture on their global dynamics. In addition, we show that both IGP models can have multiple interior equilibria under certain parameters range. These analytical results indicate that IGP model with generalist predator has “top down” regulation by comparing to IGP model with specialist predator. Our analysis and numerical simulations suggest that: (1) Both IGP models can have multiple attractors with complicated dynamical patterns; (2) Only IGP model with specialist predator can have both boundary attractor and interior attractor, i.e., whether the system has the extinction of one species or the coexistence of three species depending on initial conditions; (3) IGP model with generalist predator is prone to have coexistence of three species.     

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.     

The earliest evidence of holometabolan insect pupation in conifer wood

Background

The pre-Jurassic record of terrestrial wood borings is poorly resolved, despite body fossil evidence of insect diversification among xylophilic clades starting in the late Paleozoic. Detailed analysis of borings in petrified wood provides direct evidence of wood utilization by invertebrate animals, which typically comprises feeding behaviors.

Methodology/Principal Findings

We describe a U-shaped boring in petrified wood from the Late Triassic Chinle Formation of southern Utah that demonstrates a strong linkage between insect ontogeny and conifer wood resources. Xylokrypta durossi new ichnogenus and ichnospecies is a large excavation in wood that is backfilled with partially digested xylem, creating a secluded chamber. The tracemaker exited the chamber by way of a small vertical shaft. This sequence of behaviors is most consistent with the entrance of a larva followed by pupal quiescence and adult emergence — hallmarks of holometabolous insect ontogeny. Among the known body fossil record of Triassic insects, cupedid beetles (Coleoptera: Archostemata) are deemed the most plausible tracemakers of Xylokrypta, based on their body size and modern xylobiotic lifestyle.

Conclusions/Significance

This oldest record of pupation in fossil wood provides an alternative interpretation to borings once regarded as evidence for Triassic bees. Instead Xylokrypta suggests that early archostematan beetles were leaders in exploiting wood substrates well before modern clades of xylophages arose in the late Mesozoic.     

Borings in Quartzite Surf Boulders from the Upper Cambrian Basal Deadwood Formation,Black Hills of South Dakota

Shallow, semi-spherical borings occur in clusters with densities of 1–3.5/cm² in quartzite boulders and in vein quartz from the localized basal conglomerate of the Cambrian-Ordovician Deadwood Formation in the east-central Black Hills of South Dakota. Some borings are superimposed on primary but enigmatic semi-circular structures, 2.5–5 cm in diameter, which are soft-sediment trace fossils formed prior to lithification. The macroborings are the first to be recorded from quartzite and vein quartz. Host boulders were eroded from near vertically dipping Paleoproterozoic quartzites from several different stratigraphic units. The thin boulder conglomerate grades were abruptly changed to sandstone through a layer ≤2 m thick in the Marjuman transgression (regionally correlated to the Cedarina dakotaensis trilobite zone). This transgression occurred prior to the start of the globally recognized Upper Cambrian Paibian Stage. Physically similar rocky-shore settings are widely known from quartzite islands of Cambrian age in Wisconsin, Middle Ordovician age on Ontario's Manitoulin Island, Ordovician-Silurian age in Manitoba, and Devonian age in western Australia. Erosion of quartzite surf boulders of equal or larger size occurred in all those regions, but the Black Hills of South Dakota is the only region where borings in quartzite are documented.     

Does an opportunistic predator preferentially attack nonvigilant prey?

The dilution effect as an antipredation behaviour is the main theoretical reason for grouping in animals and states that all individuals in a group have an equal risk of being predated if equally spaced from each other and the predator. Stalking predators, however, increase their chance of attack success by preferentially targeting nonvigilant individuals, potentially making relative vigilance rates in a group relatively important in determining predation compared with the dilution effect. Many predators, however, attack opportunistically without stalking, when targeting of nonvigilant individuals may be less likely, so that the dilution effect will then be a relatively more important antipredation reason for grouping. We tested whether an opportunistically hunting predator, the sparrowhawk, Accipiter nisus, preferentially attacked vigilant or feeding prey models presented in pairs. We found that sparrowhawks attacked vigilant and feeding mounts at similar frequencies. Our results suggest that individuals should prioritize maximizing group size or individual vigilance dependent on the type of predator from which they are at risk. When the most likely predator is a stalker, individuals should aim to have the highest vigilance levels in a group, and there may be relatively little selective advantage to being in the largest group. In contrast, if the most likely predator is an opportunist, then individuals should simply aim to be in the largest group and can also spend more time foraging without compromising predation risk. For most natural systems this will mean a trade-off between the two strategies dependent on the frequency of attack of each predator type. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.     

中国湘南早侏罗世双壳类壳体内的遗迹化石--兼论遗迹化石属Talpina Hagenow, 1840

论文描述的双壳类壳体内的钻孔遗迹组合发现于湖南省东南部宜章县下坪附近的心田门组上部(早侏罗世,早辛涅缪尔期)浅海沉积地层中。这个遗迹化石群中,帚虫类动物制造的钻孔Talpinahunanensisichnosp.nov.(新遗迹化石种)明显地占据了大部分,偶尔共生的其它钻孔包括:Rogerellaichnosp.(蔓足亚纲尖胸类的遗迹)、Calcideletrixichnosp.(可能是藻类的遗迹)、藻或菌造的遗迹(暂定为"Mycelites"ichnosp.)、以及一种可能被海绵动物制造的未定钻孔。生物侵蚀者侵害了海底上较大的双壳类壳体。钻孔保存为天然印模(石核),它们是文石质壳体溶解后留下的空间中的充填物。此外,文中还讨论了遗迹化石属TalpinaHagenow,1840,尤其讨论了有关遗迹种颇为复杂的鉴别问题,同时校正了该遗迹属的属征。     

Farbmuster auf jurassischen Belemniten-Rostren

Colour patterns have been observed in belemnites of the genus Megateuthis from the middle Jurassic of North and South Germany. They are present only in adult specimens, appearing first on the dorsal and then on the ventral side, and, finally, on the flanks. This suggests that the rostrum at this stage was covered only by a thin (translucent) layer of soft tissue. More probably, the rostrum was temporarily uncovered, as indicated by the presence of borings of fungi or algae on several rostra. The regular distribution of these borings suggests pre-mortem infestation of the belemnites. Fossil bacteria have also been found within the individual colour patterns.     

Taphonomy of Logs Bored with Teredolites longissimus Kelly and Bromley in the Danian (Lower Paleocene) of West Greenland

The boring Teredolites longissimus Kelly and Bromley is recorded infesting silicified logs in the Fossil Wood Member, Kangilia Formation (Lower Danian, Paleocene) of West Greenland. There are two morphologies of T. longissimus, probably representing two different episodes of invasion of the wood. The initial borings of Morphology 1 are large, deformed to an elliptical section, and confined to a direction of boring parallel to sub-parallel to the wood grain. Morphology 2 borings are less common, narrower and shorter, sinuous, markedly crosscut the grain of the wood and, where preserved intact, retain their circular cross-section. They were a later infestation than Morphology 1. Following the infestation by Morphology 1, the logs were deformed, probably by weight of overburden, and then reworked, when some of them were infested by Morphology 2 producers. Final burial was followed by brittle collapse of some Morphology 2 borings and lithification before further deformation could occur. The occurrence of T. longissimus in deeper water deposits close to a land mass with a steep slope indicates that transport in the marine realm was minimal before waterlogging.