Microborings in mollusk shells, Bay of Safaga, Egypt: Morphometry and ichnology

Boring patterns of microbial endoliths were analyzed in bivalve shells collected from different environments in the Northern Bay of Safaga on the Red Sea, Egypt. A total of 20 ichnotaxa have been recognized within 64 samples and evaluated on the basis of 250 scanning electron micrographs. The analysis of microorganisms in shells from the same depth profile has been performed separately. Morphological properties of selected ichnotaxa were studied in detail, morphometric parameters established, measured, and statistically evaluated. This method was applied to determine the variability of dominant traces in view of their ichnotaxonomic treatment. Ichnogenus Rhopalia was discussed and a new ichnospecies Rhopalia spinosa was described. Formal change of name for cyanobacterial traces of Hyella and Solentia from Fasciculus to Fascichnus, and for the traces of the green alga Ostreobium from Reticulina to Ichnoreticulina were proposed on the ground of the priority rights in compliance with rules of the International Code of Zoological Nomenclature.     

THE FISH AND AMPHIBIAN SWIMMING TRACES UNDICHNA AND LUNICHNIUM, WITH EXAMPLES FROM THE LOWER PERMIAN OF NEW MEXICO, USA

Abstract: The ichnotaxonomy and stratigraphic, geographic and environmental distribution of fish (Undichna) and amphibian (Lunichnium) swimming traces are reviewed. The ichnospecies of Undichna consist of various combinations of sinusoidal waves of differing complexity. Some of the more complex ichnospecies are made up of elements of the simpler forms, and morphological subset relationships between them are presented. Such subset series represent potential taphoseries relationships (i.e. preservational variants that reflect, for example, undertrails), or series of minor behavioural variations. Such a system can be used to highlight that different ichnospecies occurring at a locality may be taphonomic or minor behavioural variants of each other. Caution should, therefore, be exercised before erecting new ichnospecies on the basis of limited material if its morphology is a subset of an existing ichnospecies. However, the naming of such simpler ichnospecies is valid if they represent a recurrent morphology, and it is valid to erect new ichnospecies whose morphology is not a subset of an existing ichnospecies. Specimens that demonstrate intergradation between ichnotaxa can be used to justify their synonomy. Ichnotaxonomic revisions reduce the number of ichnospecies in Undichna from 14 to nine. U. radnicensis, a highly variable ichnospecies, is synonymized with U. britannica on the basis of material from China that demonstrates they can intergrade. U. prava is a partial U. tricosta, which falls within the minimum diagnosis of U. simplicitas. U. gosiutensis is regarded as a subjective junior synonym of U. quina. U. westerbergensis, originally attributed to a ‘crossopterygian’ fish performing a tetrapod‐like gait, is reassigned as a distinct ichnospecies within Lunichnium because it demonstrates the same morphology, representing similar behaviour, albeit by a different producer. L. anceps and L. gracile are synonymized with L. rotterodium. New specimens of U. bina and L. rotterodium are also described from the Lower Permian Robledo Mountains Formation of southern New Mexico, USA.     

The Ichnogenus Kouphichnium and Related Xiphosuran Traces from the Steven C. Minkin Paleozoic Footprint Site (Union Chapel Mine), Alabama,USA: Ichnotaxonomic and Paleoenvironmental Implications

Abstract

The ichnogenus Kouphichnium and associated ichnofossils attributed to xiphosuran activity are here re-examined from samples collected from the Pennsylvanian-age Steven C. Minkin Fossil Site at the Union Chapel Mine, in Walker County, Alabama, USA. The large sample size offers an unique opportunity to evaluate some Kouphichnium ichnospecies. Thus, the morphological variability resulting from the taphonomic, ethological, taxonomic variability and underprint fallout are evaluated using this large sample set. Three morphotypes have been segregated from the material previously assigned to K. aspodon discovered at this and adjacent sites. Ichnospecies of Kouphichnium identified at the Union Chapel Mine site include: K. lithographicum, K. aspodon and two new ichnospecies (K. atkinsoni and K. minkinensis). Additionally, Kouphichnium-like traces that are associated with “jumper” traces have been previously misinterpreted as Kouphichnium and Selenichnites, respectively, and are excluded from this study, leaving them in open nomenclature as they will be the subject of a subsequent publication. We here redescribe the holotype of K. aspodon and designate lectoparatypes to better define the ichnospecies. New trace makers for some Kouphichnium ichnospecies are hypothesized, in contrast to the traditional xiphosuran attribution.     

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.     

The ichnogenus Palmiraichnus Roselli for fossil bee cells

The ichnogenus Palmiraichnus was erected by Roselli in 1987 to include his formerly described ich‐nospecies castellanosi, and a new one, minor, herein considered a junior synonym of the former. Shape, lining and closure clearly show that these traces are fossil bee cells. Celliforma, the other ichnogenus for non‐clustered fossil bee cells, includes two ichnospecies, bedfordi (=septata) (n. syn.) and pinturensis, which share with Palmiraichnus castellanosi the presence of antechambers and hardened cell walls. This combination of features is characteristic of certain groups of short‐tongued bees and justifies the inclusion of these ichnospecies in Palmiraichnus. Microscopic structures of cell walls and closures are adopted here for ichnotaxonomic purposes and for comparing fossil with extant cells revealing that they are useful ichnotaxobases with respect to the problematic fossil bee cell ichnotaxonomy. The micro‐structure of the cell walls and closures of the three ichnospecies of Palmiraichnus are compared. The presence of antechambers and particularly of the thickened cell rears suggest a relationship between Palmiraichnus castellanosi and Uruguay, an ichno‐taxon also attributable to bees. However, in Uruguay ichnospecies. cells are grouped in well‐defined clusters, whereas in Palmiraichnus castellanosi, cells occur isolated in paleosols, and no intermediate morphologies are known between the two ichnogenera. P. castellanosi, the focus of our study, is also compared with oxaeid cells revealing similarities in size and wall microstructure and differences in microstructure of closures and cell rears.     

First Early Jurassic Ornithischian and theropod footprint assemblage and a new ichnotaxon Shenmuichnus wangi ichnosp. nov. from Yunnan Province,southwestern China

The new ichnospecies, Shenmuichnus wangi ichnosp. nov., is the first evidence for the presence of large ornithischians in the Early Jurassic of Yunnan Province, whereas the known skeletal record documents small species only. Until now Shenmuichnus was known from a single locality in Shaanxi Province by the ichnospecies Shenmuichnus youngteilhardorum. Compared with the latter, Shenmuichnus wangi is larger and shows a different trackway configuration, particularly in the relative position of manus and pes imprints. Palecologically, the occurrence of Shenmuichnus wangi in a red bed facies indicates the preference of distinctive environments of trackmakers of both ichnospecies, questioning former hypotheses of exclusivity of ornithischians in more humid climates. By abundance both skeletons and footprints of ornithischians suggest their role as a minor component in Early Jurassic saurischian dominated dinosaur faunas in this region.     

Lacertoid Footprints of the Upper Permian Arenaria di Val Gardena Formation (Northern Italy)

Lacertoid footprints are the largest component of the Upper Permian Arenaria di Val Gardena Formation ichnofauna that contains hundreds of specimens mostly referred to the ichnogenus Rhynchosauroides Maidwell 1911. In this paper, we analyzed unpublished material and re-examined the Rhynchosauroides footprints of that ichnofauna, in particular the figured specimens. Analysis of Rhynchosauroides and its type ichnospecies R. rectipes Maidwell 1911 was first necessary. This preliminary investigation highlighted several problems, including ichnospecies named on the basis of poorly preserved material and in some cases significantly different from the type ichnospecies.

The study allowed for recognition of three ichnotaxa referred to Rhynchosauroides: Rhynchosauroides pallinii Conti et al., 1977, Rhynchosauroides isp.1 and Rhynchosauroides isp.2 and Ganasauripus ladinus igen. et isp. nov. Other material previously referred to Rhynchosauroides is herein regarded as unclassifiable, in the light of present ichnological knowledge and procedures.     

The Recognition of Ophiomorpha irregulaire on the Basis of Pellet Morphology: Restudy of Material from the Type Locality

Serial grinding of a specimen of Ophiomorpha irregulaire collected from Coal Creek Canyon in the Book Cliffs of Utah, USA—the type locality of the ichnospecies—allows us to assess the validity of pellet morphology as an ichnotaxobase for species-level identification. The importance of O. irregulaire stems from its abundance in petroliferous shallow marine strata of post-Paleozoic age. Our three-dimensional reconstruction of the burrow wall of O. irregulaire demonstrates for the first time that it is composed of sand-cored pellets. The typically spiky nature of the pellets in cross-section is inferred to result from differential compaction of the sand core and pelletal lining. Rupturing of the thin outer clay coating of the sand pellets is considered to produce the distinctively attenuated, spiky outer surface to the burrow wall of this ichnospecies. This study demonstrates the utility of pellet morphology in species-level classification of Ophiomorpha and lends support to O. irregulaire being a cosmopolitan ichnospecies present in post-Paleozoic strata. It also demonstrates the need to ensure that ichnotaxonomic identifications are based on three-dimensional knowledge of the trace fossil in question.     

Review of Japanese Dinosaur Track Localities: Implications for Ichnotaxonomy,Paleogeography and Stratigraphic Correlation

Nine dinosaur ichnospecies from the Lower Jurassic to Upper Cretaceous of Japan, including two that are new, are described herein. The new ichnotaxa are Asianopodus pulvinicalx ichnogen. et ichnosp. nov. and Schizograllator otariensis ichnosp. nov. The Japanese ichnotaxa are allied to Lower Jurassic ichnospecies in South China, North America, Western Europe and South Africa, and Upper Jurassic to Lower Cretaceous ichnospecies from Southeast and East Asia. This suggests they were part of a global ichnofauna before continental drift began in the Middle Jurassic, leading to the development of a more endemic dinosaur fauna in the Cretaceous. At least two assemblages, an ornithopod-gracile-toed theropod-dominated community, in northeastern Asia, and a robust theropod- and sauropod-dominated community in the southern part of the continent, existed in the Cretaceous. This parallels North American dinosaur distribution patterns in the Cretaceous and seems to be a reflection of paleolatitudinal controls.     

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.     

Carboniferous Psammichnites: Systematic Re-Evaluation,Taphonomy and Autecology

The ichnogenus Psammichnites Torell 1870 includes a wide variety of predominantly horizontal, sinuous to looped, backfilled traces, characterized by a distinctive median dorsal structure. Though commonly preserved in full relief on upper bedding surfaces, some ichnospecies of Psammichnites may be preserved in negative hyporelief. Psammichnites records the feeding activities of a subsurface animal using a siphon-like device. Several ichnogenera reflect this general behavioral pattern, including Plagiogmus Roedel 1929 and the Carboniferous ichnogenera Olivellites Fenton and Fenton 1937a and Aulichnites Fenton and Fenton 1937b. Based on analysis of specimens from the United States, Spain, and the United Kingdom, three Carboniferous ichnospecies of Psammichnites are reviewed in this paper: P. plummeri (Fenton and Fenton, 1937a), P. grumula (Romano and Meléndez 1979), and P. implexus (Rindsberg 1994). Psammichnites plummeri is the most common Carboniferous ichnospecies and is characterized by a relatively straight, continuous dorsal ridge/groove, fine transverse ridges, larger size range, and non-looping geometric pattern. It represents a grazing trace of deposit feeders. Psammichnites grumula differs from the other ichnospecies of Psammichnites by having median dorsal holes or protruding mounds. The presence of mounds or holes in P. grumula suggests a siphon that was regularly connected to the sediment-water interface. This ichnospecies is interpreted as produced by a deposit feeder using the siphon for respiration or as a device for a chemosymbiotic strategy. Psammichnites implexus is characterized by its consistently smaller size range, subtle backfill structure, and tendency to scribble. Although displaying similarities with Dictyodora scotica, P. implexus is a very shallow-tier, grazing trace. Changes in behavioral pattern, preservational style, and bedform morphology suggest a complex interplay of ecological and taphonomic controls in Carboniferous tidal-flat Psammichnites. A first distributional pattern consists of guided meandering specimens preserved in ripple troughs, probably reflecting food-searching of buried organic matter concentrated in troughs. A second is recorded by concentration of Psammichnites on ripple crests and slopes. In some cases, the course is almost straight to slightly sinuous and closely follows topographic highs, suggesting a direct control of bedform morphology on trace pattern. Occurrences of Carboniferous Psammichnites most likely represent an opportunistic strategy in marginal-marine settings. Analysis of Carboniferous Psammichnites indicates the presence of a siphon-like device in the producer and reestablishes the possibility of a molluscan tracemaker.     

Large Carnivore Footprints from the Late Pleistocene of Argentina

The aim of the present contribution is to describe large felipedid footprints from a new ichnological site from the Late Pleistocene of Buenos Aires Province, Argentina. The prints are referred as the new ichnospecies Felipeda miramarensis nov. ichnosp. Based on size and morphology, this new ichnotaxon may have been produced by the large machairodontine felid Smilodon populator. Track analysis indicates that the producer of the tracks had fully retractile claws, a plantigrade feet, and lacked strong supination capabilities on pes and manus. The size and depth differences between manus and pes prints indicate that the producer had notably robust anterior limbs. If correctly assigned, the new ichnospecies reinforces the idea that Smilodon was an ambush predatory mammal.     

Ichnologic Note A New Ichnospecies of Cardioichnus from the Cretaceous (Albian) of New Mexico

A new ichnospecies of Cardioichnus, Cardioichnus biloba isp. nov., is described and documented from shallow marine strata of Cretaceous (Late Albian) age in southern New Mexico, USA. It is a heart-shaped resting trace which, unlike all other ichnospecies of Cardioichnus, is epichnial and increases in width to a bilobate trace, often with a raised anterior terminus. Only the posterior end of this trace is associated with Bichordites burrows, and the anterior end is bilobate and undisturbed. This represents a resting position which differs from other known species of this cubichnium, which are hypichnial and subquadrate in symmetry. The tracemaker is a spatangoid echinoid, most likely Heteraster d'Orbigny, 1853.     

New ichnospecies of Linichnus Jacobsen & Bromley, 2009

Abstract

A new ichnospecies, Linichnus bromleyi, is described on bone substrate as the result of a very likely predator/scavenger interaction. L. bromleyi consists of a single groove with a non-serrated edge. This new ichnotaxon is compared with L. serratus which was defined as a single elongate serrate-edged groove. L. bromleyi has been identified over the surface of bones of marine mammals (in particular, cetaceans and pinnipeds) from two Pliocene outcrops of Andalusia, southern Spain. The new ichnospecies can be clearly related with a trophic interaction between sharks and marine mammals (mainly whales, dolphins and seals).     

The New Ichnogenus Flagrichnus—A Paleoenvironmental Indicator for Cold-Water Settings?

The new marine microboring ichnogenus Flagrichnus is described from Pleistocene mollusc shell beds of Rhodes (Greece) and the Costa Brava (Spain). Recent equivalents are studied in detail from the cold-temperate setting of the Swedish Kosterfjord area. Two ichnospecies are recognized: Flagrichnus profundus consists solely of a deeply penetrating gallery with a basal swelling while Flagrichnus baiulus comprises a single to multiple sack-shaped cavity at the base of a branching filamentous gallery that is penetrating deeply into the substrate. The traces are distributed from the euphotic zone down to aphotic depths suggesting (chemo)heterotrophic microendoliths and more specifically marine fungi as the trace makers. For Flagrichnus profundus, the thraustochytrid fungus Schizochytrium is recognized as its trace maker; for Flagrichnus baiulus, no definite candidate has been identified. While Flagrichnus profundus is distributed from tropical to arctic waters, Flagrichnus baiulus is only known from fossil and Recent non-tropical settings suggesting an applicability of this ichnospecies as an indicator for low paleotemperatures.     

Silurian craniide brachiopods from Gotland

Seven craniide brachiopod genera are described from the Silurian (Wenlock–Ludlow) of Gotland, including one new genus and five new species. The new genus and species Thulecrania septicostata is unique among Silurian craniides as it possesses solid spines. The new species Lepidocrania multilamellosa is the first Silurian record of this poorly known Permian genus. The problematic North American Propatella Grubbs, 1939 , was originally described as a gastropod, but the new species Propatella palmaria from Gotland shows that it is a craniide with sutured hollow spines of a type not previously recorded from Silurian craniides. The dorsal valves of the new species Valdiviathyris? bicornis are remarkably similar to those of the type species and represent the first possible Palaeozoic record of this poorly known extant craniide. This first systematic study of craniide brachiopods from the Silurian of Gotland shows that the diversity is relatively high as compared to other known Silurian craniide faunas, but a more thorough comparison is not possible due to the lack of data from most parts of the world. The new data from Gotland support the view that the craniides were not affected by the end‐Ordovician extinction.     

The ichnofamily Celliformidae for Celliforma and allied ichnogenera

This contribution undertakes a comprehensive revision, lacking until now, of all the ichnotaxa attributed to fossil bee cells, one of the most common traces in paleosols. These ichnotaxa are morphologically related to Celliforma and consequently grouped herein in the new ichnofamily Celliformidae. Two new ichnogenera are formulated, Cellical‐ichnns igen. nov., for several ichnospecies previously included in Celliforma, and a new ichnospecies, C. chubutensis isp. nov., belonging to this ichnofamily, and Brownichnus igen. nov., for the ichnospecies favosites, which is not morphologically related to Celliforma. The ichnogenera included in this ichnofamily involve traces in which Celliforma, the most simple trace of the group, is part of their structure. Paltniraichnus is akin to Celliforma with antechambers and discrete walls. Uruguay, Corim‐batichnus and Rosellichnus are clusters composed of adjacent rows of Celliforma, or Palmiraichnus‐like cells. Ellipsoideichnus and Cellicalichnus are different arrangements of Celliforma‐like cells attached to tunnels.

Celliformidae are based on the morphology of the traces, all of them comprising cells, groups of cells and cells attached to tunnels. As such, its component ichnotaxa are based exclusively on morphological ichnotaxobases, which, in turn, were evaluated and selected with respect to the nest architecture of the trace makers, the bees. This paper analyzes some procedures and clues (i.e. behavioral homologies) that may be used to select the proper taxobases to erect ichnotaxa when the identity of the trace makers is known, as in this study case.