Amphibian Body Impressions from the Mississippian of Pennsylvania,USA

Temnocorpichnus isaacleai is a new ichnogenus and ichnospecies of temnospondyl amphibian body impression from the Mississippian (Visean) Mauch Chunk Formation of eastern Pennsylvania, United States. The shovel-shaped head, robust limbs, relatively short trunk and smooth integument diagnose the ichnotaxon and readily distinguish it from Hermundurichnus and Sauropleura, the only other named body impressions of Paleozoic tetrapods. Temnocorpichnus is a compound ichnogenus in which the footprint ichnogenus Batrachichnus is a behaviorally distinct component. The temnospondyl identity of Temnocorpichnus adds to the sparse and earliest records of temnspondyls, which are of Visean age. The smooth integument of the ichnogenus does not support the presence of ventral scales or armor in the earliest temnospondyls, but body proportions of the Mauch Chunk body impressions indicate a relatively terrestrial temnospondyl not matched by any taxon now known from bones. Three closely associated impressions of Temnocorpichnus on a single bedding plane suggest some sort of gregarious behavior in Mississippian temnospondyls and may support speculation that internal fertilization and associated courtship behavior evolved independently in one group of amphibians more than 300 million years ago.     

A New Ichnospecies of Arthrophycus from the Late Carboniferous (Pennsylvanian) of Michigan,USA

A unique ichnofossil assemblage from Pennsylvanian-age sandstones near Eaton Rapids, Michigan, USA, comprises straight-to-curved traces preserved in convex hyporelief, with the transverse ridges and median grove associated with the ichnogenus Arthrophycus. The Michigan traces show some branching or pseudo-branching (also known from other Arthrophycus specimens) and are among the smallest structures (millimeters in diameter) attributed to this ichnogenus. The orientation of the Michigan Arthrophycus burrows is distinct from other ichnospecies of this taxon in the pronounced co-planar alignment of the burrows, as opposed to the multiplanar, interweaving, “bundled” nature typical of the ichnogenus. On this basis we assign the Michigan specimens to a new ichnospecies of Arthrophycus. The “paralleling behavior” of the new taxon may reflect a strategy of the tracemaker to avoid previous burrows and reflect differences in resource availability or current energy. This new taxon supports previous records of the occurrence of this ichnogenus in Upper Carboniferous strata.     

Trace fossils of the arthropod Camptophyllia from the Westphalian (Carboniferous) rocks of Lancashire, UK and their palaeoenvironmental context

Two arthropod trace fossils are described and analysed from the Carboniferous Lower Westphalian (C. communis and basal A. modiolaris chronozones) coal-bearing strata of Lancashire. The biserial trackway Diplichnites triassicus consists of five overlapping en echelon sets of 7–9 tracks preserved as epichnia and hypichnia in lacustrine siltstones. The trackway suggests subaqueous in-phase walking by a multi-segmented producer with a body length of 35–40 mm, width 17–22 mm, and 7–9 appendages. Curved, clustered, or laterally repeated, hypichnial lobes with transverse striations on the base of ripple cross-laminated sandstone are identified as Rusophycus versans. This trace fossil is interpreted as shallow resting or furrowing burrows of a homopodous arthropod, 30–60 mm long, 15–30 mm wide, and probably the same kind of arthropod as produced D. triassicus.A review of contemporary arthropod body fossils from Lagerstätten in Lancashire favours the onisciform, or Arthropleura like arthropod Camptophyllia as a potential producer of both of these trace fossils in a lacustrine palaeoenvironment.This study integrates the analysis of sediments, trace fossils and body fossils for reconstructing the arthropod biota and ecology in Westphalian lacustrine and crevasse splay fluvial palaeoenvironments.     

A Foraminiferal Parasite on the Sea Urchin Echinocorys: Ichnological Evidence from the Late Cretaceous (Lower Maastrichtian,Northern Germany)

Archaeonassa is a poorly known ichnogenus, originally described from the Cambrian of North America, which is a member of the Scolicia ’group’, but can be regarded as distinct from Scolicia. Archaeonassa was originally based on modern material, and therefore falls into a taxonomic grey zone concerning its availability, but as A. fossulata was erected on fossil material the ichnogenus is here retained. Although exhibiting a degree of morphological plasticity in its style of ornament, Archaeonassa is considered to be monospecific, with only the ichnospecies A. fossulata recognised. The ichnogenus appears to be restricted to the Paleozoic, although similar material is well documented from modern environments. Thus the ichnogenus is expected to be geographically and stratigraphically more widespread than it otherwise appears. Archaeonassa is a valuable tool for environmental reconstruction within the Paleozoic as it occurs within intertidal deposits. The ichnogenus is interpreted chiefly as the work of gastropods, although in many cases may have been produced by trilobites or even echinoids, and represents the exogenic expression of either surface or shallow subsurface locomotion.     

Insights on the Behavior of Late Paleozoic Aquatic Crustaceans (Pygocephalomorpha?): Compound Trace Fossils from Western Argentina

We describe intergradations between the arthropod repichnial trace fossils Diplichnites gouldi (Gevers et al., 1971), Dendroidichnites Demathieu et al., 1992, and Monomorphichnus Crimes, 1970 with two distinct cubichnial traces, Gluckstadtella cooperi Savage, 1971 and Huilmuichnus santracruzensis new ichnogenus and ichnospecies. These compound ichnofossils are part of high ichnodiversity arthropod-dominated trace fossil associations from the Late Carboniferous Agua Escondida Formation of the San Rafael Basin (Argentina). The complex ichnotaxonomy of Diplichnites and the ichnospecies D. gouldi are discussed and some possible working solutions are proposed. The most likely producer of the compound trace fossils are the Pygocephalomorpha, a group of Late Paleozoic crustaceans that are recorded in nearby basins from Uruguay and Brazil. The compound trace fossils are used to infer a number of complex behaviors of pygocephalomorphs in a subaqueous setting, including caridoid (tail flip) or more primitive escape reactions, landing followed by resting and transient resting during normal walking. In addition, different gaits can be inferred from well-preserved D. gouldi, which can be correlated with contrasting trackway morphology.     

A new sauropodomorph ichnogenus from the Lower Jurassic of Sichuan,China fills a gap in the track record

A dinosaur tracksite in the Lower Jurassic Ziliujing Formation of Sichuan Province, China consists of a spectacular sub-vertical exposure, with multiple track-bearing levels and trackways showing parallel and bimodal orientations. Based on well-preserved material, the new ichnogenus and ichnospecies, Liujianpus shunan ichnogen. nov. ichnosp. nov. is erected to accommodate distinctive sauropodomorph trackways occurring in this assemblage. Liujianpus has a unique combination of features, some relating to the early Jurassic basal sauropodomorph (prosauropod in traditional usage) ichnogenus Otozoum, others to the sauropod ichnogenus Brontopodus. Despite such a mix of basal sauropodomorph- and sauropod-like features, the trackmaker of Liujianpus is likely a basal sauropodomorph. This identification is consistent with the occurrence of basal sauropodomorph skeletons from geographically and chronologically close localities. The other distinct morphotype from the tracksite is linked to a sauropod trackmaker. As such, the ichnofauna consisting of two distinct foot morphotypes reflects the diversity of sauropodomorph dinosaurs in the Early Jurassic of Asia.     

Large Pelycosaur Footprints from the Lower Pennsylvanian of Alabama,USA

Relatively large tetrapod footprints from Lower Pennsylvanian strata of the Pottsville Formation in the Warrior basin coal field of Alabama were previously assigned to the ichnogenus Attenosaurus. However, three morphotypes are present, and we assign these tracks to Attenosaurus, a new ichnogenus (Alabamasauripus) and Dimetropus. Most of these tracks apparently were made by large pelycosaurs, for which there is no bone record older than Late Pennsylvanian.     

Probable Relationships between the Lower Jurassic Crocodilomorph Trackways Batrachopus and Selenichnus: Evidence and Implications Based on New Finds from the St. George Area Southwestern Utah

Some small and medium-sized crocodylomorph footprints are described from the Late Jurassic (Kimmeridgian) coastal and deltaic units of the northern Spain (Asturias). There are at least four footprint morphotypes. Three of them, with well preserved trackways, are included in the ichnogenus Crocodylopodus (Crocodylopodus isp. and Crocodylopodus meijidei); the fourth one, documented by some isolated large footprints, is referable to the ichnogenus Hatcherichnus. This ichnoassociation confirms the presence of small crocodilians in palaeoenvironments apparently dominated by dinosaurs. The presence of Hatcherichnus seems to confirm the affinity between the Iberian and North American ichnofaunas.     

Crocodile swim tracks from the latest Cretaceous of Europe

Recently discovered evidence of tracks in the continental beds of the Late Cretaceous Tremp Formation in the southern Pyrenees (NE Iberian Peninsula) has been identified as scratch marks made by buoyant crocodiles. The tracks are preserved in two distinct environments and substrates (marly limestones originating in a littoral mud flat and fine‐grained sandstones deposited in fluvial settings). Most of the crocodylian traces are ascribed to ichnogenus Characichnos, whereas a single plantigrade pes track is assigned to ichnogenus cf. Crocodylopodus. The crocodylian swim traces (Characichnos ichnofacies) found in the early and late Maastrichtian co‐occur with Brontopodus ichnofacies attributable to terrestrial tetrapods (titanosaur sauropods, cf. Brontopodus ichnogenus; and hadrosaurid ornithopods, Hadrosauropodus ichnogenus). Analysis of the tracks allows the interpretation of palaeoenvironmental settings and track production. Thus, in lagoonal environments, swim tracks of crocodylians were produced during the rise of the water level in successive tide cycles; in fluvial settings, the swim traces of crocodylians were produced within the channel at the low‐water stage. To date, there are no reports of Late Cretaceous crocodylian tracks in Europe, and the studied evidence represents the first and youngest track record of the group in the latest part of the Cretaceous (C29r) in this continent and probably in the world.     

Lipid content of terrestrial arthropods in relation to body size,phylogeny, ontogeny and sex

Energy storage in arthropods has important implications for survival and reproduction. The lipid content of 276 species of adult arthropods with wet mass in the range 0.2–6.13 g is determined to assess how lipid mass scales with body mass. The relative contribution of lipids to total body mass is investigated with respect to phylogeny, ontogeny and sex. The lipid content of adult insects, arachnids, and arthropods in general shows an isometric scaling relationship with respect to body mass (M) (M_{arthropod lipid} = ?1.09 ×M_dry^1.01 and M_{arthropod lipid} = ?1.00 ×M_lean^0.98). However, lipid allocation varies between arthropod taxa, as well as with sex and developmental stage within arthropod taxa. Female insects and arachnids generally have higher lipid contents than males, and larval holometabolous insects and juvenile arachnids have higher lipid contents than adults.     

Behaviour of Dinosaurian Track-Makers in the Winton Formation (Cretaceous,Albian–Cenomanian) at Lark Quarry,Western Queensland,Australia: Running or Swimming?

A remarkable assemblage of dinosaur trackways in the Winton Formation (Albian–Cenomanian) at Lark Quarry, in western Queensland, Australia, has long been regarded as evidence of a stampede involving small theropods, whose tracks were classified in the ichnogenus Skartopus, and small ornithopods, whose tracks represented a second ichnogenus, Wintonopus. However, one recently-published study has claimed that existing interpretation of Lark Quarry is incorrect: it maintains that all the track-makers were ornithopods, that the ichnogenus Skartopus is a variant form of Wintonopus, and that most of the Lark Quarry track-makers were not running but more probably swimming downstream in a current of water. Those iconoclastic claims are investigated here and shown to be untenable. They derive from overgeneralized interpretation of the Lark Quarry track assemblage and invoke ad hoc auxiliary hypotheses which are either untestable or demonstrably incorrect. Closer inspection of the evidence underpinning those claims corroborates the existing interpretation of Lark Quarry as the site of a dinosaurian stampede and confirms the validity of the original distinction between theropod tracks (Skartopus) and ornithopod tracks (Wintonopus).     

The first record of Anomoepus tracks from the Middle Jurassic of Henan Province,Central China

Small, gracile mostly tridactyl tracks from the Middle Jurassic of Henan Province represent the first example of the ichnogenus Anomoepus from this region. They represent a growing number of reports (at least eight) of this ichnogenus from the Jurassic of China. In conjunction with Changpeipus and Eubrontes, they appear characteristic of known global footprint biochrons. Anomoepus indicates the presence of ornithischian dinosaurs that are often scarce or unknown from skeletal remains in coeval deposits. When first discovered, these tracks were informally referred to as bird tracks. This interpretation reflects convergence between small Jurassic Anomoepus and avian theropod tracks that are hitherto known only from the Cretaceous and the Cenozoic. However, most Anomoepus are larger and more robust than any hitherto known Mesozoic avian tracks.     

The insect trace fossil Tonganoxichnus from the middle Pennsylvanian of Indiana: Paleobiologic and paleoenvironmental implications

The ichnogenus Tonganoxichnus, produced by one or more monuran insect taxa, is now recorded from the Middle Pennsylvanian Mansfield Formation of Indiana. Tonganoxichnus is a resting trace that has three important implications. First, it represents a recurrent behavioral pattern in Upper Carboniferous to Lower Permian marginal marine environments of North America. Second, it provides finely resolved anatomical information for axial and appendicular body structures and behaviors that are difficult to determine from body‐fossil material alone. Third, integrated sedimentologic and ichnologic observations indicate that the Tonganoxichnus assemblage, inclusive of other ichnotaxa, is common in tidal rhythmites that were developed under freshwater conditions, probably in the innermost part of estuarine systems, close to or at the fluvioestuarine transition.     

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.     

First Thyreophoran Type Tracks from the Middle Jurassic Chuanjie Formation of Yunnan Province,China

Two tridactyl footprints from the Chuanjie Formation (Middle Jurassic) of Yunnan Province, China are morphological characteristics of thyreophoran tracks. They show some similarities to Shenmuichnus, known from the Early Jurassic strata of both Shaanxi and Yunnan provinces, but are somewhat larger, thereby resembling the ichnogenus Stegopodus. Based on their general morphology and size being congruent with this ichnogenus, they are tentatively assigned here to cf. Stegopodus. This is the fourth report of large ornithischian (probably thyreophoran) tracks from the Lower-Middle Jurassic of China that indicates relatively large trackmakers that were likely to be taxonomically distinct from much smaller and gracile Anomoepus trackmakers, also of ornithischian affinity. The larger tracks indicate a hitherto unreported abundance, size range and diversity of track types attributed to this group. The parallel orientation of the two best preserved trackways may indicate gregariousness.     

Enalikter is not an annelid: homology,autapomorphies and the interpretation of problematic fossils

A megacheiran arthropod, Enalikter aphson, was recently described by Siveter et al. (2014) from the mid‐Silurian (late Wenlock) of Herefordshire. Previously, megacheirans had only been recognized from the Cambrian. Struck et al. (2015) considered the body plan of Enalikter to be incompatible with this affinity, arguing that many of the arthropod features were either not present or misinterpreted. Instead, they compared Enalikter to polychaete annelids, identifying characters from numerous polychaete lineages which they considered to be present in Enalikter. A reply to this critique by Siveter et al. (2015) reaffirmed arthropod affinities for Enalikter by presenting additional evidence for key arthropod features, such as arthropodized appendages. Here, we augment Siveter et al. by critically addressing the putative annelid characters of Enalikter presented by Struck et al. and additionally explore the morphological and phylogenetic implications of their hypothesis. We conclude that similarities between Enalikter and polychaetes are superficial and that character combinations proposed by Struck et al. are not present in any annelid, living or extinct. This taxon highlights the importance of using a phylogenetic framework for interpreting fossils that present unusual morphologies, such that proposed shared characters are hypotheses of homology rather than merely phenotypic similarities. Crucially, we argue that autapomorphic characters of subgroups of large taxa (like families or classes within phyla) should not be used to diagnose problematic fossils.     

Development and evaluation of an enzyme-linked immunosorbent assay to detect Pieris rapae remains in guts of arthropod predators

An enzyme-linked immunosorbent assay (ELISA) was developed to detect remains of Pieris rapae L. (Lepidoptera: Pieridae) immature stages in the guts of field collected arthropod predators. The assay can be used to help ascertain the relative importance of arthropod predator species in suppressing P. rapae in cabbage, Brassica oleracea var. capitata L. The ELISA is sensitive to all immature stages of P. rapae, although first and fifth instars can be detected more readily than eggs or pupae and third instars showed intermediate detectability. Assays on whole body homogenates of predators readily detected predation on P. rapae first instars by all seven of the predator species tested, although response generally declined with increasing predator size. Together the results show that the P. rapae ELISA possesses a sufficiently high level of sensitivity and specificity to be a useful tool in helping to elucidate the roles of arthropod predator species in reducing populations of P. rapae in cabbage.     

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.