Arthrophycus as a mimic of crinoid column impressions in the lower Silurian of central Kentucky,USA

Unusual specimens of an ichnofossil, herein ascribed to Arthrophycus, are described from siltstone tempestites in the lower Silurian (Llandovery, Telychian) Estill Shale near Irvine, Estill County, central Kentucky, USA. The specimens display closely spaced, uncurved transverse annulations, but lack the medial ridge, rectangular cross‐sectional shape and dense branching typical of common Silurian Arthrophycus ichnospecies. This combination of features, coupled with the straightness of burrows and alignment parallel to inorganic tool marks, closely approximates the appearance of moulds of partially articulated crinoid columns (= pluricolumnals) that were impressed into firm mud and cast by storm‐deposited sediment. That these structures do not represent the external moulds of body fossils is suggested by minor width variations along the long axis of specimens, possible bioglyphs and the total absence of skeletal material, including isolated crinoid columnals, in this and other tempestites from this interval. More convincingly, the presence of very sparse branching indicates a trace fossil origin for these structures. These fossils are best interpreted as the products of preferential mining of inorganically generated tool marks by deposit feeders, owing to exposure of more organic‐rich muds by erosional scouring. Thus, this occurrence represents a rare instance in which trace fossils display a strong alignment parallel to inorganic sedimentary structures.     

Trace fossils in middle to late triassic fluvial redbeds,pranhita‐godavari valley,south India

Several morphological varieties of trace fossils abound in Middle and Late Triassic fluvial redbeds in the Pranhita‐Godavari Valley, south India, including Skolithos, Palaeophycus, Taenidium, escape burrows, and a type of trace very similar to ‘small stuffed burrows’ from the Triassic of Greenland. Burrow morphology was influenced by local hydrodynamic conditions. The distribution of burrows was facies controlled; some forms are restricted to channel deposits whereas others occur only in floodplains. Vertical dwelling burrows (Skolithos) occur in both channel and floodplain deposits. Horizontal structures representing deposit feeding (Taenidium) are confined to nondepositional surfaces within parallel‐laminated sandstones having parting lineations that represent catastrophically emplaced sand‐sheets in channels and proximal floodplains. Vertical escape burrows are confined to what were slowly but continually accreting parallel‐laminated sands of channel bars. Horizontal dwelling burrows (Palaeophycus) and ‘small stuffed burrows’ are virtually restricted to the smaller sandsheets of floodplain drainage systems.

The burrow assemblages do not occur as recurrent associations throughout the redbed sequence, and variations in different stratigraphic levels seem to be controlled by minor differences within a broadly similar environment. The entire assemblage has components of both the Scoyenia and Rusophycus ichnocoenoses reported from East Greenland but may be considered as the Scoyenia ichnofacies characteristic of redbeds deposited in extensive floodplains dissected by small streams, even though no Scoyenia individuals are present.     

Three-Dimensional Morphological and Permeability Modelling of Diplocraterion

Three-dimensional mini-permeametry of Diplocraterion burrows from the Middle Jurassic Scarborough Formation of North Yorkshire, UK, reveals some of the relationships between ichnofossils and reservoir quality in conventional siliciclastic petroleum reservoir facies. This work focuses on the effect that the various Diplocraterion burrowing behaviours have on the permeability characteristics of the near-burrow environment. Samples were studied using serial grinding and mini-permeametry. This is the first documented attempt at combining these techniques to create an integrated understanding of the inter relationship of trace fossil morphology and reservoir quality in three dimensions. Palaeobiological insights arising from the three-dimensional modelling include evidence for resuspension feeding, deposit feeding, sediment cleaning and “collapse-cone feeding.” Higher than expected permeability values (>150 mD) were measured from the sandy shafts of Diplocraterion and adjacent areas. Zones of enhanced permeability associated with the inter-shaft area are inferred to result from sediment cleaning, and grain-size selective deposit feeding. Bioturbation by Diplocraterion is found to improve reservoir quality by sediment cleaning and creation of highly permeable vertical conduits in a reservoir facies with otherwise low permeability.     

Silurian trace fossils in carbonate turbidites from the Alexander arc of southeastern Alaska

Early to Late Silurian (Llandovery‐Ludlow) body and trace fossils from the Heceta Formation of southeastern Alaska are preserved in the oldest widespread carbonates in the Alexander terrane. These fossils represent the earliest benthos to inhabit diverse shallow and deep subtidal environments in the region and are important indicators of early stages in benthic community development within the evolving Alexander arc. The ichnofossils are particularly significant because they add to a small but growing body of knowledge about trace fossils in deep‐water carbonates of Paleozoic age.

Carbonate turbidites that originated along a deep marine slope within the arc yield a low‐diversity suite of trace fossils consisting of five distinct biogenic forms. Simple burrows (Planolites, two forms), ramifying tunnels (Chondrites), and tiny cylindrical burrows (?Chondrites) represent the feeding activities (fodinichnia) of pre‐turbidite animals that burrowed in the lime mud before the influx of coarser sediment deposited by turbidity currents. These trace fossils are associated locally with cross‐cutting burrows created as domichnia (Palaeophycus). Rarer hypichnial burrows and endichnial traces were created by post‐turbidite animals that fed soon after the deposition of coarse detritus from turbidity flows.

Trace fossils in these deposits reflect much lower diversity levels than in Paleozoic siliciclastic turbidites. This difference may represent unfavorable environmental conditions for infaunas, differential preservation, or significant paleogeographic isolation of the Alexander terrane during the Silurian. Greater utilization of trace fossils in terrane analysis may help to resolve this issue and provide new data for reconstructing the paleogeography of circum‐Pacific terranes.     

Conichnus Männil 1966, not conichnus Myannil 1966

Vertically curved, unbranched, strongly lined burrows are prominent features of muddy outer‐shelf/offshore deposits of Cretaceous to Pleistocene age. Such burrows are usually called Terebellina, but use of this ichnogenus should be avoided for two reasons: 1) the type species of Terebellina is a large agglutinated foraminiferid referable to Bathysiphon, not a trace fossil; and 2) most trace fossils identified with Terebellina are either examples of Palaeophycus (in cases where burrows are essentially horizontal and occur individually) or Schaubcylindrichnus (where burrows are oblique to stratification and occur in aggregations or loose clusters). The replacement name S. freyi is proposed for this latter category of burrow.

These light‐colored, grain‐lined burrows are conspicuous structures in the dark‐colored, outer‐shelf mudrocks of the lower Pleistocene Rio Dell Formation. The burrows are part of a typical “distal Cruziana ichnofacies”; association of structures, including Teichichnus, Planolites, Palaeophycus, and Chondrites, occurring in thoroughly bioturbated background sediments. Schaubcylindrichnus freyi in the Rio Dell is interpreted as the dwelling structure of a gregarious, endobenthic organism, comparable to the lined burrows produced by the deposit‐feeding polychaete Clymenella.     

Furrows and firmgrounds: evidence for predation and implications for Palaeozoic substrate evolution in Rusophycus burrows from the Silurian of New York

Tarhan, L.G., Jensen, S. & Droser, M.L. 2011: Furrows and firmgrounds: evidence for predation and implications for Palaeozoic substrate evolution in Rusophycus burrows from the Silurian of New York. Lethaia, Vol. 45, pp. 329–341. The Silurian Herkimer Formation of east‐central New York contains abundant, exceptionally preserved composite Rusophycus‐Teichichnus burrows. We suggest that the most likely interpretation of these composite trace fossils is as structures formed by trilobites entering the sediment in search of prey. Parallel alignment of the paired traces, asymmetrical configuration of the Teichichnus along the longitudinal axis of the associated Rusophycus, depth correlation and deformation of the Teichichnus all suggest that this relationship was predatory. In addition, sectioned material indicates that these Rusophycus may have been open at the sediment‐water interface, while the crisp preservation of both Rusophycus and Teichichnus, along with the preservation of such delicate morphological details as scratch marks, suggests that the sediment must have been relatively firm at the time the traces were formed. The formation and preservation of Rusophycus in cohesive sediments located very close to the sediment‐water interface hold important implications for the manner in which we consider Palaeozoic substrates and their temporal and spatial evolution. Moreover, these findings demonstrate that the morphology and taphonomy of ichnological associations may, in the context of sedimentological relationships, prove a powerful proxy for tracking substrate conditions through both space and time. □firmgrounds, New York, predation, Rusophycus, substrate, taphonomy.     

Triassic trace fossils from lacustrine shoreline deposits of the Passaic formation,douglassville, pennsylvania

Lake‐margin deposits of the Late Triassic Passaic Formation, Douglassville, Pennsylvania, have yielded a moderate variety of trace fossils. The greatest diversity and abundance of trace fossils occurs on the sole of a thin gray claystone overlain and underlain by gray siltstones. Specimens of Cochlichnus anguineus, Helminthoidichnites tenuis, Helminthopsis hieroglyphica, Treptichnus pollardi, and paired trails reflect simple, unspecialized, horizontal grazing as well as feeding traces under very shallow water lacustrine conditions. The lack of commonly associated Scoyenia burrows in these deposits may possibly be related to the degree of maturation of the organic debris available. The traces, as well as a lack of meniscate burrows, compare favorably to the Mermia ichnofacies, except Passaic deposits exhibit evidence of subaerial exposure. This unit most likely records a lacustrine expansion whereby grazing trails were emplaced under fully subaqueous conditions. Subsequent shallowing and desiccation, followed by sediment influx during rainstorms, favored preservation of these traces.

In contrast, the Scoyenia ichnofacies consists of feeding burrows of Scoyenia gracilis and Spongeliomorpha milfordensis within reddish brown siltstones and mud‐stones (redbed sequence). The Scoyenia ichnofacies records limited exploitation, by opportunistic infaunal deposit feeders, of lake‐margin nutrients carried in during occasional rainstorms that punctuated otherwise extended periods of aridity.     

Ichnology of an ephemeral lacustrine/alluvial plain system: Jurassic East Berlin Formation,Hartford Basin,USA

A trace fossil assemblage from the Lower Jurassic East Berlin Formation of the Newark Supergroup, Hartford Basin, New England, USA, includes: Scoyenia gracilis, Skolithos ichnosp., Palaeophycus striatus, Planolites montanus, Fuersichnus ichnosp., fusiform burrows, pelleted material, an escape structure, and large burrows.

This assemblage is assigned to the Scoyenia ichnofacies. Specific lebensspuren are not limited to specific lithofacies; instead, their initial distribution seems to have been influenced principally by water availability within an ephemeral lacustrine/alluvial plain system. Other factors in distribution may have included amounts of organic matter, patterns of sedimentation, sediment grain size, biotic factors (settling from invertebrate drift, competition), and additional abiotic factors (wind deflation, waves, currents, desiccation, soft‐sediment deformation, evaporite formation, pedoturbation).

Extreme environmental conditions within the original depositional setting strongly influenced the availability of water which, in turn, strongly influenced the paleoecology of burrowing invertebrates in this nonmarine system.     

The morphology and mode of formation of <Emphasis Type="Italic">Neoeione</Emphasis> igen. nov. from the Carboniferous of northern England

Samples of the burrows studied herein, originally introduced as Eione moniliformis Tate, have been collected from the heterolithic sandstone and siltstone beds of a coastal outcrop near the town of Howick, Northumberland, UK. These interbedded and hummocky cross-stratified beds, belonging to the Stainmore Formation, are Carboniferous and represent offshore marine sediment deposited between the fair wave base and storm wave base. As the name Eione was used previously to describe a genus of gastropod, Eione Tate is a junior homonym of Eione Rafinesque. Therefore, a new ichnogenus was established, Parataenidium Buckman, to accommodate burrows composed of two distinct levels, which included E. moniliformis. Using three-dimensional morphological modeling techniques, this study aims to: (1) clarify the current taxonomic status of Parataenidium moniliformis; and (2) provide an updated model for its construction. The transfer of E. moniliformis to Parataenidium was problematic, primarily because it is not composed of two distinct levels, but, instead, is composed of backfilled reniform sediment packages. These packages were created through a multistage process whereby the organism maintained a small open cavity from which it fed. Accordingly, a new ichnogenus, Neoeione, is proposed to accommodate the material originally described by Tate.     

A review of the biology, ecology and potential use of the common ragworm Hediste diversicolor (O.F. Müller) (Annelida: Polychaeta)

The common ragworm Hediste diversicolor inhabits the shallow marine and brackish waters in the North Temperate Zone of the Atlantic. This infaunal species builds U or Y-shaped burrows in the soft sediments. H. diversicolor is gonochoristic and remains atokous throughout its life. Due to the facility of harvesting of individuals by hand and their maintenance, this species represents a good biological material for experimental studies. In consequence, the common ragworm served as a model for endocrine control of reproduction in Polychaeta and played a fundamental role in the study of immune defence in Polychaeta.Life history characteristics (longevity, spawning season, feeding tactics and growth) and population dynamics (sex ratio, density and biomass) vary greatly according to geographical location of the populations. In addition, inter-population morphological, biochemical and physiological differences have been noticed in individuals from different areas and different environmental conditions in the same areas and could be related to the limited dispersal capacity of the species. H. diversicolor interacts with its environment. The construction of burrows by individuals increases the sediment–water interface. When they ventilate their burrows, individuals verticalize oxic zones into the sediment and promote microbial and meiofaunal growth alongside their burrows. Moreover, the common ragworm is a species of commercial interest because of its use as bait in recreational fishing and as food in aquaculture. In order to alleviate environmental pressure caused by excessive demand over optimal sustainable yield of bait fisheries, intensive aquaculture has been proposed. The common ragworm could be also used as biomonitor of pollution in estuarine ecosystems.     

Analysis of Mineral Segregation in Euzonus mucronata Burrow Structures: One Possible Method Used in the Construction of Ancient Macaronichnus segregates

Mineralogical segregation of sand grains distinguishes the trace fossil Macaronichnus segregatis, which is composed of a felsic burrow infill with a mafic-and mica-rich burrow mantle. This study focuses on determining the mechanism by which M. segregatis trace-makers segregated mineral grains during deposit feeding. A modern opheliid polychaete, Euzonus mucronata, from Pachena Bay, Vancouver Island (Canada), was examined to explain the activities of their ancient counterparts. Microscopic videotaping of deposit feeding allowed for collection of data on ingestion and excretion through visual grain counts of felsic, mafic, and shell components. Normalization of these grain counts to the composition of the host sediment illustrates preferential ingestion of felsic grains over mafic. Shell fragments were generally avoided and visually mantled the burrows, obscuring the paucity of mafic grains in burrow infills. The avoidance of shell fragments is potentially a function of the large grain size, angular shape, surface texture, and/or associated low nutritive value. The preferential ingestion of felsic grains is attributed to en masse feeding in felsic-rich locales identified through sediment probing. This form of mineral segregation likely reflects the specific nature of the sediment and worm population. Accordingly, en mass deposit feeding in selected felsic-rich localities is one possible mechanism used in the construction of Macaronichnus segregatis and M. segregatis-like structures.     

Stellavelum arborensis igen., sp. nov., Stellavelum uncinum igen., isp. nov. and Termitichnus namibiensis isp. nov.; New Ichnofossils from Cenozoic deposits of Namaqualand,South Africa

Three new ichnotaxa are described from Cenozoic paralic to continental sediments on the west coast of South Africa. Stellavelum arborensis and Stellavelum uncinum are assigned to a mixed Skolithos and Cruziana ichnofacies and Termitichnus namibiensis is assigned to the Termitichnus ichnofacies. An interpretation of the sedimentary record shows that the Stellavelum trace makers were probably sediment ingesting annelids that lived in dense colonies in sheltered lagoonal environments during Neogene times. The Termitichnus namibiensis trace fossils are interpreted as actively backfilled termite nests which form an early to mid‐Pleistocene palimpsest, terrestrial ichnofabric which is superimposed on the older marine sediments.     

Roller Coaster Behavior in the Cruziana Rugosa Group from Penha Garcia (Portugal): Implications for the Feeding Program of Trilobites

Trilobite burrows of the Cruziana rugosa group are common and well preserved in the Armorican Quartzite Formation (Lower to Middle Ordovician) of the Ponsul River gorge in Penha Garcia. Based on morphological and behavioral peculiarities, Cruziana beirensis is reinstated herein and included in the rugosa group. Apart from the broad range in morphology and size, Cruziana from Penha Garcia show higher behavioral diversity using food sources than has ever been documented in a single section. This behavioral diversity, mainly in circling behavior, has been analyzed using the Capacity Fractal Dimension by implementation of the box-counting theorem applied to the bedding plane. Fractal Dimension also suggests that Cruziana rouaulti can be included in the rugosa group, despite the obscurity of the scratch patterns, as products of juveniles. Circling, sinusoidal or teichichnoid behavior modifications reflect a generalist mode of sediment feeding, claimed mudtrophobacterivory, while most of the interactions with worm burrows originated previously or (mostly) later, by interpreted trilobite necrophagy and/or worm commensalism. Patchy exploitation of biomat grazing fields is inferred from Cruziana preservation styles, physical interactions with biomat-related sedimentary structures and area-limited high bioturbational indices in the explored tier.     

Paleoecology of the marine endobenthos

Endobenthic animals, which reside within the sea bottom, include stationary suspension feeders, mobile deposit feeders and both stationary and mobile carnivores. Their activities, especially with regard to dwelling, feeding, walking/crawling and resting/nesting, are recorded as trace fossils.Abundance, diversity and density of some kinds of trace fossils allow interpretation of the population strategies of the trace-makers in terms of opportunistic (r-selected) and equilibrium (K-selected) strategies. Opportunistic ichnotaxa tend to be faciesbreaking traces, which are highly localized in low-diversity, high-density trace fossil associations in rocks representing environmental extremes (e.g., variable salinities, harsh temperatures, low oxygen levels or shifting substrates). Equilibrium ichnotaxa usually are restricted to particular sedimentary facies and are characteristic of high-diversity, low-dominance trace fossil associations in sediments reflecting stable, predictable environmental conditions.The most important environmental factors influencing the composition of trace fossil assemblages in marine settings are bathymetry, substrate, oxygen and hydrodynamic energy. The four factors are closely interrelated, because as water depth increases, there is a general decrease in sediment grain size and hydrodynamic energy of the depositional environment. As depth below the water—sediment interface increases, the firmness of the sediment (due to compaction and dewatering) increases and the oxygen content of interstitial waters drops drastically.Marine ichnofacies are largely substrate-controlled. Soupgrounds are water-saturated, incompetent substrates typified by highly compressed and usually unidentifiable burrows. Softgrounds commonly contain numerous distinctive burrows and are zoned bathymetrically by the Skolithos, Cruziana, Zoophycos and Nereites Ichnofacies. Firmgrounds are characterized by stiff, compacted sediments, in which traces of the Glossifungites Ichnofacies are excavated. Hardgrounds are cemented substrates, in which bioerosion traces of the Trypanites Ichnofacies are bored. Woodgrounds are woody materials that have been exposed to the sea and bored by bivalves, which produce characteristic traces of the Teredolites Ichnofacies. Tiering of endobenthic communities is common and is related to substrate preference of the burrowers and oxygen stratification of interstitial waters.     

Evidence for targeted elasmobranch predation on thalassinidean shrimp in the Miocene Taliao Formation,NE Taiwan

Terrestrial and marine invertebrate organisms both leave records of their activities in the sediment in the form of trace fossils, at least during certain stages of their ontogeny. In contrast, trace fossils produced by vertebrate organisms are scarce, although terrestrial trace fossils provide exclusive insights into the social behaviour of their producers. In the marine realm, vertebrate trace fossils are relatively rare, difficult to identify and problematic to interpret. However, in certain settings, observations on serendipitously preserved and exposed trace fossils can shed light on the predatory behaviour of marine vertebrates. In Miocene outer shelf to nearshore sandstones of the Taliao Formation in NE Taiwan, large numbers of bowl‐shaped trace fossils can be observed. Morphology and size range (diameter typically 10–30 cm, average depth around 10 cm) of these trace fossils agree well with feeding traces of modern stingrays, and the trace fossil Piscichnus waitemata, which has been attributed to bottom feeding rays. Stingrays direct a jet of water from their mouths to excavate a bowl‐shaped pit to expose their prey. In the material filling the excavated bowl, broken pieces of two other common trace fossils, Ophiomorpha and Schaubcylindrichnus, are often found, and in a number of cases, vertical shafts of Ophiomorpha surrounded by dispersed pieces of wall material have been observed. In contrast, surrounding sediment rarely contains this kind of broken pieces of wall material. These observations clearly indicate that stingrays specifically targeted the producers of the trace fossils: thalassinoid crustaceans and worms, respectively. The targeted predation of these relatively deep burrowers furthermore suggests that the rays used their electroreceptive organs to locate the prey; as such, direct targeting of buried prey only based on olfactory senses has been shown to be ineffective in experiments with extant myliobatiform rays.     

Scolicia shirahamensis isp. nov.: a triple-corded scolicia and its ichnological implications

Abstract

Peculiar meniscate burrows with three sediment cords occur in early to middle Miocene tidal-flat deposits of southwestern Japan. Two of the cords are situated at the bottom and the other is at its center. Detailed observations of the burrow structures and comparative neoichnological studies of modern spatangoid burrows in a tidal flat revealed that the former two were true drainage tubes and the latter was fecal in origin. The trace fossil was thus assigned to the ichnogenus Scolicia. Based on these findings, a new ichnospecies Scolicia shirahamensis isp. nov. has been described here. The central sediment cord is seemingly identical to the drainage tube of the ichnogenus Bichordites, another ichnogenus that has been commonly ascribed to a fossil spatangoid burrow, similar to Scolicia. Careless ichnogeneric identification of a spatangoid burrow, based only on the central sediment cord, therefore, may produce an incorrect identification.     

LATE PRECAMBRIAN TRACE FOSSILS FROM NEW SOUTH WALES

Seven trace-fossil species are described from the upper part of the Torrowangee Group (Upper Proterozoic) of western New South Wales, and a variety of other traces are illustrated. A few forms occur in the Fowlers Gap Beds, and a more diverse and abundant fauna is recorded from the stratigraphically higher Lintiss Vale Beds. Virtually all the traces are preserved in the plane of bedding, as semi-reliefs. The named trace fossils all come from the Lintiss Vale Beds, and are as follows: Planolites ballandus sp. nov., Planolites? sp., Cochlichnus serpens sp. nov., Gordia? sp., Torrowangea rosei gen. et sp. nov., Phycodes? antecedent sp. nov., and Curvolithus? davidis sp. nov. Three of these species are regarded as feeding burrows (endogene), and are thought to represent the activity of infaunal, worm-like deposit feeders. Others may be either feeding burrows or crawling trails. There are also a few impressions which seem to be rest marks. A discussion of the significance of the trace-fossil occurrences is presented.     

ECOLOGY AND EVOLUTION OF MATING SYSTEMS OF FIDDLER CRABS (GENUS UCA)

1. General accounts of the natural history and behaviour of fiddler crabs suggest there exist two broad mating patterns in the genus. Most western and Indo-Pacific species mate on the surface of intertidal substrates near burrows females defend. The sexes associate only briefly during courtship and mating. In contrast, males of many American species court from and defend burrows to which females come for mating. Copulation occurs underground in burrows plugged at the surface; the sexes usually remain together for at least several hours. Here we summarize and contrast recent detailed field studies of the mating systems of U. pugilator, an American species, and U. vocans, a species widely distributed in the western and Indo-Pacific. We indicate how differences in the breeding ecology of these two species may account for basic differences in modes of sexual selection leading to the two broad mating patterns in the genus. 2. U. pugilator burrows in protected sandy substrates in the upper intertidal and supratidal zone. During ebb tide, nonbreeding crabs leave burrows they occupy during high tide to forage on food-rich substrates in the lower intertidal zone. Reproductively active males remain in the burrow zone where they fight for and defend burrows from which they court. Large males win most fights for burrows and tend to defend burrows high on the elevation gradient, especially during periods with relatively high tides. Females usually approach and descend the burrows of several males before choosing their mates by remaining in males' burrows. Males remain underground with their mates for 1–3 days until after they oviposit their eggs. Some males then emerge and leave their burrows while others sequester their mates in the chambers where mating and oviposition has occurred, dig new chambers and resume courtship, perhaps attracting additional females. In either case, females remain underground for approximately 2 weeks, finally emerging to release their planktonic larvae. Burrows that do not collapse due to tidal inundation or flooding by groundwater are best for breeding and usually are located relatively high on the elevation gradient. Females choose mates indirectly by preferring to breed in burrows that will remain intact while they oviposit and incubate their eggs. Large males mate more often than small males because they are better able to defend burrows at locations females prefer to breed. The mating system of U. pugilator may be classified as resource-defence polygyny. 3. U. vocans burrows in open muddy substrates in the mid- to lower intertidal zone. At a site near Chunda Bay, Australia, where the reproductive behaviour of this species has been studied in depth, both sexes feed near burrows they defend. Females tend to occupy their burrows for longer periods and move shorter distances than do males. Mating occurs on the surface near the burrows that females defend. Females accept both resident and wandering males as mates. They show no preference for mating with larger males. Female choice may be based on other male morphological or behavioural characteristics. Females oviposit their eggs either while on the surface or in their burrows. They produce relatively small clutches and are active on the surface throughout their breeding periods. Males fight both their neighbours and wandering males. Large males tend to win fights and defend burrows in areas where large females, which produce relatively many eggs, are most dense. Such areas may offer greater protection from predators than areas occupied by smaller females. Small males mate about as often as large males but may father fewer larvae. The mating system of U. vocans is resource-free and promiscuous. 4. The mating systems of U. pugilator and U. vocans differ fundamentally in that female U. pugilator require access to a specific microenvironment to breed successfully, while female U. vocans do not. We suggest this difference occurs because of contrasts in clutch sizes and the mobility and movement patterns of feeding females. Female U. pugilator produce relatively large clutches and probably experience more intense selection from factors that can cause egg loss and mortality than do U. oocans, which produce clutches of sufficiently small volume to be protected by their abdominal flaps. Hence, the range of suitable breeding environments for U. pugilator is small compared to that for U. vocans. In addition, U. pugilator burrows in areas that are relatively food-poor, leading to daily migrations to and from food-rich substrates in the lower intertidal zone, preventing female defence of an area suitable for both breeding and feeding. U. vocans, however, burrows in areas sufficiently rich to support feeding, leading to relatively low female mobility and defence of burrows that are also suitable breeding sites. 5. Adaptive radiation of the genus Uca in the Americas is manifest by trends toward smaller adult size, higher population densities, more frequent microgeographic sympatry and increased terrestriality, compared to species in the western and Indo-Pacific regions. We outline the general features of the selection mechanisms tying each of these trends to the evolution of resource—defence mating systems. Intraspecific variation in the courtship behaviour and site of mating in U. lactea and U. vocans supports our contention that resourse—defence behaviour tends to occur at high population densities. Additional data are needed to evaluate the other hypotheses critically.     

河南泌阳凹陷古近纪核桃园组遗迹化石组合及其环境意义——兼论深水湖泊遗迹相特征

南襄盆地泌阳凹陷古近纪核桃园组湖泊沉积中发育大量遗迹化石,含5个遗迹组合,形成于从滨湖到深湖的不同沉积环境中。Taenidiumserpentinum遗迹组合反映稍有覆水并周期性暴露的低能滨湖环境;Skolithos linearis遗迹组合代表湖泊中上临滨环境;Planolites montanus遗迹组合与低能浅湖环境有关;Skolithos verticalis遗迹组合形成于浅湖的远源风暴沉积中;Semirotundichnus dongyingensis遗迹组合发育在贫氧的半深湖-深湖沉积环境中。     

Turbulence,displacement, death and worms: a day in the life of a fluvial Carboniferous bivalve

Kane, I.A. 2010: Turbulence, displacement, death and worms: a day in the life of a fluvial Carboniferous bivalve. Lethaia, Vol. 43, pp. 381–395. In the Pennsylvanian Rough Rock Flags and Rough Rock of northern England trace fossils attributed to the non‐marine bivalve Carbonicola are found. Carbonicola, recorded by Lockeia and associated trace fossils, lived a semi‐infaunal lifestyle and thus were influenced by both the sediment in which they were hosted, and the currents which supplied their nutrients and oxygen. A number of palaeocurrent indictors are commonly associated with Lockeia: (1) downstream inclination of vertical burrows; (2) palaeoflow‐parallel orientation of long axes; (3) steeper scouring and higher sediment surface on the upstream side; and (4) diffuse lamination downstream of the trace or more widespread downstream erosion. Semi‐infaunal, Carbonicola bivalves were partly exposed to the prevailing flow and acted as bed defects; flow separation and acceleration enhanced flow turbulence around the bivalve leading to erosion and the development of a variably developed fan‐shaped zone of scour immediately downstream. Disturbance and destabilization of sediment in this way may have affected bivalves immediately downstream, plausibly explaining the relatively regular spacing pattern of individual or clustered Lockeia, exposed on bedding planes and revealed by nearest neighbour analyses. Bivalves that did not survive high‐energy flow events were either trapped within the sediment, or transported downstream and deposited in lower energy environments within the otherwise high‐energy deposits of the Rough Rock.