The origin and paleoecologic significance of the trace fossil Asteriacites in the Pennsylvanian of Kansas and Missouri

The trace fossil Asteriacites, recorded in Cambrian to Recent shallow- and deep-marine facies, is traditionally interpreted as the resting trace of asterozoans. Well-preserved specimens of A. lumbricalis are abundant in Pennsylvanian (Upper Carboniferous) shallow- and marginal-marine siliciclastic deposits of eastern Kansas and western Missouri. Detailed morphologic analysis of these specimens suggests that they record the activities of mobile epifaunal ophiuroids. Evidence of a brittle star (ophiuroid) producer rather than sea star (asteroid) is provided by (1) trace-fossil morphologic features reflecting the anatomy of the producer (e.g., well-differentiated central structure, slender vermiform arms) and ophiuroid burrowing technique (e.g., proximal arm expansion, arm branching), and (2) mode of occurrence (e.g., gregarious behavior, horizontal and vertical repetition). Vertical and horizontal repetition produces complex aggregates of A. lumbricalis that are interpreted either as escape structures (fugichnia) or as feeding structures, respectively. Ophiura texturata is proposed as a modern analogue for the A. lumbricalis producer, based on inferred life habit and feeding behavior. Asteriacites lumbricalis is present in two different intertidal trace-fossil assemblages. The first assemblage is characterized by high diversity and records tidal flats developed outside of embayments under normal marine conditions. The second assemblage consists of A. lumbricalis together with a few other ichnotaxa and represents a depauperate association that developed in restricted tidal flats within an embayment or estuarine setting. This challenges the conventional view of Asteriacites as a normal-marine salinity indicator. Some echinoderms, and particularly asterozoans, penetrate and inhabit modern environments of depressed salinity. The presence of Asteriacites in Pennsylvanian marginal-marine facies of Kansas and Missouri provides evidence that ophiuroids had adapted to brackish-water conditions by the late Paleozoic.     

Asteriacites lumbricalis von schlotheim 1820: Ophiuroid trace fossils from the lower triassic thaynes formation,central Utah

Several trace fossils of burrowing ophiuroids, Asteriacites lumbricalis von Schlotheim 1820, occur on slabs of thinly‐bedded silty limestone in the lower Spathian part of the Lower Triassic Thaynes Formation in central Utah. This is the first record of Asteriacites in the Lower Triassic of North America. Their occurrence adds information to studies of the recovery of marine faunas following the Permian extinctions, including the oxygénation of benthic sediments.     

Palaeoenvironmental implications of the ichnology and geochemistry of the Westbury Formation (Rhaetian), Westbury‐on‐Severn,south‐west England

Abstract: The Westbury Formation (Rhaetian) beds of Westbury Garden Cliff, Westbury‐on‐Severn, west of Gloucester, Britain, show an unusual combination of features. Both deep water and emergent characteristics are present within the sediments and the trace fossils. The ichnoassemblage consists of abundant Selenichnites, Planolites beverlyensis and Lockeia with rarer Oniscoidichnus, Chondrites, Rhizocorallium irregulare, Taenidium serpentium, an unusual form of Walcottia and Merostomichnites‐like traces. These trace fossils display an interesting relationship with the sediments: low‐energy Cruziana ichnofacies is found within high‐energy sandstones. The sandstones are interbedded with laminated mudstones, apparently deposited in deep water, but some aspects of the ichnoassemblage, preservation and sedimentation indicate shallower water. One new trace fossil, Radichnus allingtona igen. et isp. nov., closely resembles the traces of modern fiddler crabs and imply emergence, by analogy. This ichnofauna is similar to early stage disaster colonisation in recent experiments in Long Island Sound (south of Connecticut, USA) and with storm‐influenced deposits within the Cardium Formation (Seebe, Alberta, Canada). This indicates a lagoonal environment with influxes of sand and oxygen. Total organic carbon levels were found to fluctuate greatly between stratigraphic layers but remained relatively high. This implies low oxygen conditions. The abundance of sulphur (in pyrite) also supports an interpretation of anoxic conditions, and low sedimentation rates within the shale layers. A restricted shallow basin or lagoonal environment is proposed for the palaeoenvironment, with fluctuating oxygen influencing diversity.     

Complex behavioural patterns and ethological analysis of the trace fossil Zoophycos: evidence from the Lower Devonian of South China

The trace fossil Zoophycos is abundant in the shallow‐marine deposits (tempestites) of the Lower Devonian (Emsian) Yangmaba Formation in Ganxi of Sichuan, South China. It often occurs as part of complex trace fossils that comprise different integrated elements: scratch traces, simple to complex spreiten structures with marginal tubes (Zoophycos) and vertical tunnels. The complex Zoophycos burrows consist of spreiten with a marginal tube, preserved as convex hyporeliefs on the sole of an erosion surface. The exquisite, complex spreiten are interpreted to have been formed by deposit‐feeding behaviour, where the animal constructed the trace upwards without leaving faeces in the spreiten. The width of the marginal tube in different whorls is almost constant. The scratches are observed on the wall of the marginal tubes. The Zoophycos intergrades with Spongeliomorpha and Chondrites and was later cut by vertical shafts. All these features together indicate that the Zoophycos‐maker might have been a vermiform polychaete instead of a predator such as a decapod crustacean (Spongeliomorpha producer). Based on stratigraphical and ichnological features, the complex trace fossils resulted from the complex activity of different opportunistic organisms (r‐strategist) that quickly occupied and thrived within the quiet, nutrient‐rich environment after storm events.     

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.     

An escape response behaviour in the brittle star ophiopteris papillosa (echinodermata: Ophiuroidea)

This study describes a behaviour of the brittle star Ophiopteris papillosa and tests two hypotheses concerning its possible function. During this behaviour an individual rapidly raises all five arms above its disk as the brittle star falls through the water column. The final position, with the five arms held tightly together above the disk, is termed the tulip position. Individuals in the tulip position fell faster, thus reaching the bottom sooner, than individuals not in the tulip position when tested in the laboratory. Individuals in the tulip position were consumed less frequently than individuals not in the tulip position when tested in the field. I conclude that the tulip position increases the likelihood of a brittle star reaching the protective cover of the bottom and escaping fatal predation.     

Regional variation in fish predation intensity: a historical perspective in the Gulf of Maine

Summary Regional variation in the intensity of fish predation on tethered brittle stars and crabs was measured at 30–33 m depths in the rocky subtidal zone at seven sites representing coastal and offshore regions of the Gulf of Maine, USA. Analysis of covariance comparing the slopes of brittle star survivorship curves followed by multiple comparisons tests revealed five groupings of sites, with significantly greater predation rates in the two offshore than in the three coastal groups. Brittle stars tethered at the three offshore sites were consumed primarily by cod, Gadus morhua, with 60–100% prey mortality occuring in 2.5 h. In striking contrast, only 6–28% of brittle star prey was consumed in the same amount of time at the four coastal sites, which were dominated by cunner, Tautogolabrus adspersus. In several coastal trials, a majority of brittle star prey remained after 24 h. The pattern of higher predation offshore held for rock crabs as well with only 2.7% of tethered crabs consumed (n=36) at coastal sites versus 57.8% of crabs (n=64) consumed at offshore sites. Another important predatory fish, the wolffish, Anarhichas lupus, consumed more tethered crabs than brittle stars. Videos and time-lapse movies indicated that cod and wolffish were significantly more abundant at offshore than at coastal sites. Three hundred years of fishing pressure in New England has severely depleted stocks of at least one important benthic predator, the cod, in coastal waters. We speculate that this human-induced predator removal has lowered predation pressure on crabs and other large mobile epibenthos in deep coastal communities. Transect data indicate that coastal sites with few cod support significantly higher densities of crabs than offshore sites with abundant cod.     

Bichordites monastiriensis from the Pleistocene of southeast Jamaica

Bichordites monastiriensis Plaziat and Mahmoudi 1988, produced by burrowing spatangoid echinoids, is figured and described from shallow marine siliciclastic strata of the Pleistocene Old Pera Beds (Coastal Group) of southeast Jamaica. Its occurrence there represents only the second formal recording of this monospecific ichno‐taxon and extends its geographic range from the Mediterranean region to the Caribbean. The spatangoid species that produced this trace fossil is unknown, but, unlike examples from other areas, it is unlikely to be Echinocardium.     

Feeding biology of the Antarctic brittle star Ophionotus victoriae (Echinodermata: Ophiuroidea)

Summary The diet and feeding behavior of the brittle star Ophionotus victoriae were studied in 1981–1983. Frequency-of-occurrence and points methods were used to examine stomach contents of 2475 specimens from 19 stations representing seven general localities along the Antarctic Peninsula and South Sandwich Islands. Results of the two methods were in agreement and indicated this species is an opportunistic generalist with high diet plasticity. The most abundant component of the stomach contents was sediment but the diet derived from at least 13 phyla. Crustaceans, notably euphausiids and amphipods, were most important at some locations. Diet composition varied significantly between locations and years, but did not differ among brittle star size groups. Stomach content data and laboratory studies indicate that O. victoriae utilizes a variety of feeding methods but not suspension feeding.     

Parataenidium,a new taenidium‐like ichnogenus from the carboniferous of Ireland

The new ichnogenus Parataenidium is erected for backfilled tubular trace fossils that can appear superficially similar to Taenidium, but are divided horizontally into two distinct levels. Two ichnospecies are recognised: Parataenidium mullaghmorensis isp. nov. and Parataenidium moniliformis (Tate 1859). The latter ichnospecies is transferred from Eione Tate 1859, which is a junior homonym of Eione Rafinesque 1814, and therefore unavailable for Tate's ichnotaxon. The ichnogenus is an important component of late Paleozoic shallow‐water siliciclastic sediments, and can be considered as a “guide”; indicator for the Carboniferous.     

Abundance,biomass and spatial distribution pattern of brittle stars (Echinodermata: Ophiuroidea) on the Kolbeinsey Ridge north of Iceland

Species composition, abundance, biomass and spatial distribution pattern of brittle star assemblages of the Kolbeinsey Ridge, north of Iceland, were investigated by analysing seafloor photographs and epibenthic sled catches. Sampling was conducted in July 1992 along a 34 km long cross-ridge transect at 67°55N in depths ranging from 830 to 1100 m. Five brittle star species were found. Only one species, Ophiocten gracilis, occurred with densities of more than 1 ind m^–2. Both ridge slopes were characterized by soft bottom habitats, the ophiuroid distribution, however, revealed differences between slopes. Generally, brittle star densities were significantly higher on the eastern slope. There, the highest mean abundance per station of O. gracilis was 497 ind m^–2. Except for one station, a distinct patchiness on the 100 m-scale was obvious in the spatial distribution of O. gracilis. Small settling stages with disc diameters <1 mm accounted for up to 98% of the population of O. gracilis near the ridge top, but their abundance share decreased with depth to only 6% on the eastern slope foot. Biomasses of O. gracilis were extrapolated by combining abundances estimated from photographic counts with a size-weight relationship and size frequencies established from sled catches. The highest value (120 mg AFDW m^–2) was found on the deep eastern slope where large O. gracilis with disc diameters >4 mm were most abundant. Our findings reflect a cross-ridge gradient in terms of hydrographic regime and, hence, probably pattern of food supply for the benthos.     

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.     

Brittle star fauna (Echinodermata: Ophiuroidea) of the arctic northwestern Barents sea: composition,abundance, biomass and spatial distribution

 Epibenthic brittle star assemblages were investigated on the northwestern Barents Sea shelf between 81° and 77°N in July 1991. At 9 drift stations in water depths between 80 and 360 m, series of 35–71 photographs, each depicting about 1 m² of the seabed, were taken along transects of about 150- to 300-m length to assess abundances and spatial distribution patterns of adult brittle stars (disc diameter ≥1 mm). Biomass values were derived by combining abundances with size-weight relationships and size frequencies established using specimens from trawl catches. Six brittle star species were identified on the seabed images. Ophiocten sericeum was the most abundant species on shallow shelf banks (≤100 m). Up to 2,800 individuals were counted on a single photograph; median abundances per station ranged from 32 to 524 ind.m^-2 and biomass from 0.3 to 5.0 g ash-free dry weight (AFDW) m^-2. The spatial distribution along the transects (i.e. on the 100-m scale) was, however, extremely patchy. Disc diameters of O. sericeum ranged between 1.6 mm and 15.4 mm. In deeper shelf habitats (>150 m), O. sericeum was rare or absent, and Ophiacantha bidentata dominated the brittle star fauna with median densities and biomasses of 2–49 ind.m^-2 and 0.07–1.9 g AFDW m^-2, respectively. Its disc diameters ranged from 2.9 to 14.4 mm. The other species (Ophiura sarsi, Ophiopholis aculeata, Ophioscolex glacialis, Ophiopleura borealis) occurred in distinctly lower numbers. Our findings provide further evidence that brittle stars dominate epibenthic communities on Arctic shelves and locally reach very high abundances. Dense beds of Ophiocten sericeum seem to be a general phenomenon on high-Arctic shallow shelf banks. Received: 30 March 1995/Accepted: 30 June 1995     

Commentary

A new ichnotaxon is recognized in the Perkasie Member of the Passaic Formation (Upper Triassic) at the Smith Clark Quarry, Milford, New Jersey. Spongeliomorpha milfordensis is a small trace fossil possessing sharply incised partial wall striations which form a distinctive angle with the axis of the burrow. It is construed as a deposit‐feeding structure possibly formed by aquatic insect larvae along the margin of a lake.     

Asteroid and Ophiuroid Trace Fossils from the Lower Cretaceous of Chile

Abundant asterozoan trace fossils in the Lower Cretaceous Apeleg Formation of southern Chile were produced by the infilling of traces made by asteroids and ophiuroids on a muddy surface. Most are preserved as hypichnial ridges formed as casts on the bottom of fine-grained sandstone laminae. Star-shaped Asteriacites lumbricalis are interpreted as the infillings of shallow excavations made by asteroids. Hook-shaped and sinuous ridges of the newly defined ichnogenus and ichnospecies Ophioichnus aysenensis are interpreted as the casts of imprints made by the walking action of the arms of ophiuroids. Deposition probably took place in an offshore tide-swept marine shelf environment. In life the animals formed an assemblage equivalent to echinoderm aggregations on the modern sea floor.     

Commentary

Recently rediscovered type material referred to Muensteria Stern‐berg confirms that in its original (paleobotanical) definition this genus is too heterogenous to be useful in ichnotaxonomy. It contains plant remains, a coprolite string, and trace fossils. Some corrections concerning appearance of the analyzed specimens are introduced.     

Precambrian–Cambrian boundary interval occurrence and form of the enigmatic tubular body fossil Shaanxilithes ningqiangensis from the Lesser Himalaya of India

The affinity of the Ediacaran fossil Shaanxilithes ningqiangensis and putatively related forms has long been enigmatic; over the past few decades, interpretations ranging from trace fossils to algae to metazoans of uncertain phylogenetic placement have been proposed. Combined morphological and geochemical evidence from a new occurrence of S. ningqiangensis in the Krol and Tal groups of the Lesser Himalaya of India indicates that S. ningqiangensis is not a trace fossil, but rather an organic‐walled tubular body fossil of unknown taxonomic affinity. Specimens consist of compressed organic cylindrical structures, characterized by extended, overlapping or fragmented iterated units. Where specimens intersect, overlapping rather than branching or intraplanar crossing is observed. Lithologic comparisons and sequence stratigraphic data all suggest a late Ediacaran age for the uppermost Krol Group and basalmost Tal Group. By extending the biogeographical distribution of S. ningqiangensis, hitherto confined to the Ediacaran of China and potentially Siberia, to the Precambrian–Cambrian boundary interval of India, this new occurrence of S. ningqiangensis expands the biostratigraphic utility of this enigmatic fossil to the inter‐regional and intercontinental scale. Moreover, study of these new and exceptionally preserved samples may help to significantly constrain the long‐debated problem of Shaanxilithes' affinity, elucidating its ‘problematic’ status and shedding new light upon the ecology and taphonomy of one of the most significant intervals in early life history.