Holes in Bones: Ichnotaxonomy of Bone Borings

Bone is a substrate for bioerosion at equal rank with xylic and lithic substrates. Accordingly, borings in bone have to be identified in an analogous way to other ichnogenera coined for one type of substrate. In due course, the new ichnogenera Osteichnus n. igen. and Clavichnus n. igen. are established within the new ichnofamily Osteichnidae. Gastrochaenolites and Trypanites are here restricted to lithic substrates, and Asthenopodichnium only occurs in xylic substrates. Only with this approach, ichnotaxobases of trace fossils in bone are identical to those in other hard substrates. Cuniculichnus variabilis n. igen. n. isp. is introduced for variably shaped pits to tunnels bored into bone by beetle (arguably dermestid) larvae; its ethological character is close to a pupichnion.     

EARLY DEVONIAN BIOEROSION IN THE RADE DE BREST, ARMORICAN MASSIF, FRANCE

Abstract:  Abundant and diverse bioerosion trace fossils in the Lower Devonian of the Rade de Brest, France, were produced by boring ctenostomate bryozoans, polychaete annelids and algae. Two unidentifiable traces of encrusting sclerobionts are also considered. Two new taxa are described: Pinaceocladichnus cristatus ichnosp. nov. and Ichnogutta erectus ichnogen. et ichnosp. nov. Both are attributed to the work of boring bryozoans. Sedimentary condensation during intervals of sea-level rise seems to have had an important influence in promoting infestation by organisms, the activities of which are now preserved as trace fossils. A zonation of bioerosion trace fossils within the offshore zone, in the Lower Devonian of the Armorican Massif, is also suggested.     

Bioerosion trace fossils on bones of the Cretaceous South American theropod Buitreraptor gonzalezorum Makovicky,Apesteguía and Agnolín, 2005 (Deinonychosauria)

The ichnological record provides valuable information on the lifestyle, behaviour, and other palaeobiological and palaeoecological aspects of the biota. Here, we describe an interesting case of bioerosion trace fossils in bones of Buitreraptor gonzalezorum Makovicky, Apesteguía and Agnolín, 2005, a deinonychosaurian theropod from the fossiliferous locality of La Buitrera, Río Negro, Patagonia, Argentina. The trace fossils are morphologically diverse and preserved in a great percentage of the skeleton, including the jaw, vertebrae and limbs. Four main groups of trace fossils have been informally named as Parallel-Edge Furrows, Overlapped Grooves, Punctures and Lined. Parallel-Edge Furrows are in turn subdivided into four subgroups: isolated furrows, parallel pairs, opposed pairs and a combination of parallel and opposed pairs. The bioerosion trace fossils were probably generated by scavenging activities, and the semi-articulated preservation of the skeleton and the small size of each individual trace indicate small-sized tracemakers. Mammals are the main candidates although some traces may have been generated by crocodyliforms and insects such as dermestids and termites. This evidence provides additional information about palaeoenvironmental conditions, taphonomic processes, taxonomic diversity and ecological relationships that characterised this part of northern Patagonia at Early Cretaceous times.     

Über Lebensspuren aus dem eozänen Belluno-Flysch (Nord-Italien)

An inventory of trace fossils from the Eocene Bellunese Flysch is given for 10 localities. The 19 ichnogenera belong to the ethological groups Fodinichnia and Pascichnia. Taxonomic and nomenclatory remarks on some ichnogenera are made. Comparisons with other Flysch trace fossil assemblages prove a typical Flysch ichnocoenosis.     

Names for trace fossils: a uniform approach

The taxonomic treatment of trace fossils needs a uniform approach, independent of the ethologic groups concerned. To this aim, trace fossils are rigorously defined with regard to biological taxa and physical sedimentary structures. Potential ichnotaxobases are evaluated, with morphology resulting as the most important criterion. For trace fossils related to bioerosion and herbivory, substrate plays a key role, as well as composition for coprolites. Size, producer, age, facies and preservation are rejected as ichnotaxobases. Separate names for undertracks and other poorly preserved material should gradually be replaced by ichnotaxa based on well-preserved specimens. Recent traces may be identified using established trace fossil taxa but new names can only be based on fossil material, even if the distinction between recent and fossil may frequently remain arbitrary. It is stressed that ichnotaxa must not be incorporated into biological taxa in systematics. Composite trace fossil structures (complex structures made by the combined activity of two or more species) have no ichnotaxonomic standing but compound traces (complex structures made by one individual tracemaker) may be named separately under certain provisions. The following emendations are proposed to the International Code of Zoological Nomenclature: The term 'work of an animal' should be deleted from the code, and ichnotaxa should be based solely on trace fossils as defined herein.     

Temperate bioerosion: ichnodiversity and biodiversity from intertidal to bathyal depths (Azores)

In the temperate Azores carbonate factory, a substantial fraction of the calcareous skeletal components is recycled by a remarkable biodiversity of biota producing bioerosion traces (incipient trace fossils). To study this biodiversity, experimental carbonate substrates were exposed to colonisation by epilithic and endolithic organisms along a bathymetrical gradient from 0 to 500 m depth, during 1 and 2 years of exposure. The overall bioerosion ichnodiversity is very high and comprises 56 ichnotaxa and ichnoforms attributed to cyanobacteria, chlorophytes, fungi, other micro-chemotrophs, macroborers, grazers and epilithic attachment scars. In the intertidal, hydrodynamic force, partial emersion and strong temperature fluctuations lead to the lowest ichnospecies richness. This contrasts with the highest ichnodiversity found at 15 m under the most favourable environmental conditions. Towards aphotic depths, a gradual depletion in ichnodiversity is observed, most probably because of the restricted light availability and a slowdown in ichnocoenosis development. Analysis of similarity (ANOSIM), in combination with non-metrical multidimensional scaling (NMDS), was used to highlight variability in the relative abundance of traces among depths, substrate orientations and exposure times. Ichnodiversity and abundance of traces decrease significantly with depth and are higher on up-facing versus down-facing substrates, whereas differences between years were not as pronounced. This study demonstrates that statistical methods of biodiversity analysis are not per se restricted to biotaxa but may well be applied also to ichnotaxa. In the analysis of trace fossil assemblages, this approach supports the recognition of diversity patterns and their relation to environmental gradients.     

山东莱阳盆地早白垩世莱阳群的遗迹化石

山东莱阳早白垩阳群自下而上分为瓦屋夼组,林寺山组,止风庄组,水南组,龙旺庄组和曲格庄组。为一套河湖相沉积,产有较丰富,分异度较高的非海相遗迹化石和兽脚类恐龙足迹化石,共鉴定出遗迹属11个,未定属1个,其中遗迹种9个,未定种2个,遗迹化石是：Cochlichnus anguines,Diplocraterion parallelun,Helminthoidichnites tenuis,Palaeophycus tubularis,Planolites montanus ,Scolica sp.,Scoyenia gracilis,Skolithos linearis,Taenidium cameromensis,Thalassinoides sp.,恐龙足迹化石是：Paragrallator yangi,这些化石按习性可分为4类,即：居住构造, 爬行迹,觅食[迹和牧食迹,其中又以前3种为主,遗迹化石在剖面上的分布不均匀,以上部的水南组,龙旺庄,曲格庄3个组最丰富。     

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.     

New Miocene scarabeid and hymenopterous nests and early miocene (santacrucian) paleoenvironments,patagonian Argentina

Three new trace fossils are described from Miocene paleosols of southern Argentina. Celliforma pinturensis, n. ichnosp. and Celliforma rosellii, n. ichnosp. are interpreted as cells of digging bees, possibly Anthophoridae, and Coprinisphaerafrenguellii, n. ichnosp. are brood balls of dung‐beetles. Both burrowing bees and dung‐beetles are common nesters in relatively open areas, confirming previous reconstructions of the paleoenvironment of the Pinturas Formation. A brief review of scarabeid and bee fossil nests from South America is presented, and we propose that constructed nests have a higher preservation potential than excavated nests. This fact explains their more common occurrences as trace fossils in paleosols. A new ethological category, calichnia, is proposed for hymenopterous and coleopterous traces, in which adult individuals make nests exclusively for larvae.     

The bioeroded megasurface of Oura (Algarve,south Portugal): implications for the Neogene stratigraphy and tectonic evolution of southwest Iberia

The use of rocky palaeoshore bioerosion analysis as a tool to solve stratigraphic and tectonic issues is beginning to bear fruits. The occurrence of an extensive intra-Miocene marine abrasion platform in southern Portugal at Oura (Albufeira) has been identified on the basis of bioerosion trace fossils analysis. The observed ichnodiversity is rather low, with bivalve boring Gastrochaenolites being dominant. Nevertheless, the ichnoassemblage may be assigned to the Entobia ichnofacies. The palaeoichnological study of the Oura hardground confirmed the existence of an important intra-Miocene stratigraphic gap (ca. 3 Ma hiatus), represented by a razor-sharp erosional contact that separates the two main Neogene units in the Algarvian region: the lower carbonate sequence of Lagos–Portimão Formation (Langhian/Serravallian) and the upper siliciclastic sequence of the Cacela Formation (Upper Tortonian).     

Bioerosive structures from Miocene marine mobile‐substrate communities in southern Spain,and description of a new sponge boring

Abstract: Neogene palaeoshore sediments are abundantly represented along the Mediterranean coast of Iberia. An outcrop north of the Sierra Tejeda, named La Resinera, exposes concentrations of pebbles and boulders of marble, comprising an upper Miocene marine beach deposit. The high diversity of bioerosion trace fossils present in these boulders includes structures produced by polychaete annelids, demosponges, echinoids and endolithic bivalves, which indicate a shallow shoreface environment. The ichnotaxa represented are Maeandropolydora sulcans, Caulostrepsis taeniola, Entobia geometrica, Entobia ovula, Circolites kotoncensis, Gastrochaenolites torpedo, Gastrochaenolites lapidicus, Gastrochaenolites ornatus and Gastrochaenolites turbinatus. The borings are Tortonian (late Miocene) in age. Also present, and particularly abundant, are large sponge borings that have a single chamber from which radiating canals emerge. This trace fossil is designated as Entobia resinensis isp. nov.     

‘Ten Years After’—a long-term settlement and bioerosion experiment in an Arctic rhodolith bed (Mosselbukta,Svalbard)

Rhodolith beds and bioherms formed by ecosystem engineering crustose coralline algae support the northernmost centres of carbonate production, referred to as polar cold-water carbonate factories. Yet, little is known about biodiversity and recruitment of these hard-bottom communities or the bioeroders degrading them, and there is a demand for carbonate budgets to include respective rates of polar carbonate build-up and bioerosion. To address these issues, a 10-year settlement and bioerosion experiment was carried out at the Arctic Svalbard archipelago in and downslope of a rhodolith bed. The calcifiers recorded on experimental settlement tiles (56 taxa) were dominated by bryozoans, serpulids and foraminiferans. The majority of the bioerosion traces (30 ichnotaxa) were microborings, followed by attachment etchings and grazing traces. Biodiversity metrics show that calcifier diversity and bioerosion ichnodiversity are both elevated in the rhodolith bed, if compared to adjacent aphotic waters, but these differences are statistically insignificant. Accordingly, there were only low to moderate dissimilarities in the calcifier community structure and bioerosion trace assemblages between the two depth stations (46 and 127 m), substrate orientations (up- and down-facing) and substrate types (PVC and limestone), in that order of relevance. In contrast, surface coverage as well as the carbonate accretion and bioerosion rates were all significantly elevated in the rhodolith bed, reflecting higher abundance or size of calcifiers and bioerosion traces. All three measures were highest for up-facing substrates at 46 m, with a mean coverage of 78.2% (on PVC substrates), a mean accretion rate of 24.6 g m^?2  year^?1 (PVC), and a mean bioerosion rate of ?35.1 g m^?2 year^?1 (limestone). Differences in these metrics depend on the same order of factors than the community structure. Considering all limestone substrates of the two platforms, carbonate accretion and bioerosion were nearly in balance at a net rate of ?2.5 g m^?2 year^?1. A latitudinal comparison with previous settlement studies in the North Atlantic suggests that despite the harsh polar environment there is neither a depletion in the diversity of hard-bottom calcifier communities nor in the ichnodiversity of grazing traces, attachment etchings and microborings formed by organotrophs. In contrast, microborings produced by phototrophs are strongly depleted because of limitations in the availability of light (condensed photic zonation, polar night, shading by sea ice). Also, macroborings were almost absent, surprisingly. With respect to carbonate production, the Svalbard carbonate factory marks the low end of a latitudinal gradient while bioerosion rates are similar or even higher than at comparable depth or photic regime at lower latitudes, although this might not apply to shallow euphotic waters (not covered in our experiment), given the observed depletion in bioeroding microphytes and macroborers. While echinoid grazing is particularly relevant for the bioerosion in the rhodolith bed, respective rates are far lower than those reported from tropical shallow-water coral reefs. The slow pace of carbonate production but relatively high rates of bioerosion (both promoted by low carbonate supersaturation states in Arctic waters), in concert with high retention of skeletal carbonates on the seafloor and no calcite cements forming in open pore space created by microborers, suggest a low fossilisation potential for polar carbonates, such as those formed in the Mosselbukta rhodolith beds.     

Ichnology of deep‐sea fan overbank deposits of the Ganei slates (Eocene,Switzerland)— a classical flysch trace fossil locality studied first by Oswald Heer

Ganei (Switzerland) is a classical locality for trace fossils. At this site, Heer (1877) described a large number of trace fossils, several of which were new taxa. The trace fossils occur in thin‐bedded turbidites in which the basal divisions of the Bouma sequence are typically absent; the turbidites are assigned to the Ganei Slates and are Eocene in age. They are interpreted to have been deposited in an overbank environment within an upper to middle fan area distal to a channel. Two trace‐fossil associations occur: the first (I) is characterized by bulldozing organisms producing biodeformational structures, Scolica, and Nereites irregularis; the second (II) association shows a distinct tiering pattern with near‐surface graphoglyptids and a mixed layer with simple tubes such as cf. Palaeophycus and Planolites, plus patterned tubes such as Nereites cirrinalis, and Chondrites. Deeper turbidite layers were colonized by Chondrites and Gyro‐phyllites. All trace fossils show a normal size spectrum compared to previously studied trace‐fossil associations, so the degree of oxygenation probably did not influence the composition of either trace‐fossil association. Seafloor sediment was probably soft and did not affect the trace‐fossil associations. Sedimentation rate and event frequency did not change and are estimated to have been in a range of 5–10 cm/1000 years and 2–5 events per 1000 years, respectively. The composition of trace‐fossil associations I and II is therefore interpreted to have been controlled by the benthic food content being higher for trace‐fossil association I than for II.     

The changing face of the deep: Colonization of the Early Ordovician deep-sea floor, Puna, northwest Argentina

An Upper Tremadocian deep-sea ichnofauna from the Chiquero Formation of Puna, northwest Argentina, represents a link between Ediacaran and Cambrian microbial-mat dominated ecosystems and younger Ordovician deep-marine trace-fossil assemblages. This ichnofauna is preserved at the base of thin-bedded turbidites formed in the lobe fringe of a back-arc deep-sea fan. While Ediacaran–Cambrian deep-marine trace fossils are typically linked to matground grazing and feeding, microbial textures in the Chiquero Formation are rare and not associated with trace fossils. Morphologic patterns (e.g. radial trace fossils and networks) of the Chiquero ichnofauna indicate the onset of novel trophic types, recording trapping of microorganisms and bacterial farming. However, in comparison with younger Ordovician deep-sea ichnofaunas, graphoglyptids are relatively rare, poorly diverse, and geometrically simpler. This study indicates that the Early Ordovician was a pivotal point in the ecology of deep-sea infaunal communities. This Upper Tremadocian ichnofauna records the arrival of the Agronomic Revolution to the deep sea. Comparisons with slightly older and younger deep-sea ichnofaunas demonstrate that the colonization of the deep sea was a protracted process spanning the Early Paleozoic, lagging behind colonization of nearshore and offshore substrates.     

贵州台江早、中寒武世凯里组的遗迹化石

凯里组发现遗迹化石９属１４种，主要是以ＰｈｙｃｏｄｅｓｐｅｄｕｍＳｅｉｌａｃｈｅｒ为主的遗迹群落，重要的遗迹属如：Ｃｏｃｈｌｉｃｈｎｕｓ，Ｇｏｒｄｉａ，Ｍｏｎｏｍｏｒｐｈｉｃｈｎｕｓ，Ｏｌｄｈａｍｉａ，Ｐｌａｎｏｌｉｔｅｓ，Ｔｒｅｐｔｉｃｈｎｕｓ等常见于世界各地寒武纪Ｃｒｕｚｉａｎａ遗迹相。根据遗迹化石表明凯里组沉积于盐份、含氧量正常，海水能量中等的浅海软基底潮下带环境。     

Lithostratigraphy of the Lower Cambrian metaclastics and their age based on trace fossils in the Sand?kl? region, southwestern Turkey

In the Sand?kl? region of the Taurus Range of Turkey, greater than 3000 m in thickness metamorphosed siliciclastics and volcanics (Kocayayla Group) underlies the trilobite-and conodont-bearing Middle-Late Cambrian Hudai Quartzite and Çaltepe Formation.The Kocayayla Group, previously regarded as Infracambrian or Precambrian, is dated for the first time as Early Cambrian on the basis of trace fossils. Cruziana ?fasciculata, C. ?salomonis, ?Cruziana isp., ?Diplichnites isp., Monomorphichnus isp., Petalichnus isp., Rusophycus ?avalonensis, R. ?latus, Arenicolites isp., cf. Altichnus foeyni, Planolites isp., Skolithos isp., and ?Treptichnus isp. have been recognised. These trace fossils are considered Tommotian or younger in age but older than the overlying, trilobite and conodont bearing Middle Cambrian limestones of the Çaltepe Formation. The trace fossils were likely produced by trilobites, suspension feeding annelids and deposit feeding “worms”, probably polychaetes. Sections bearing abundant Skolithos represent the Skolithos ichnofacies, which is typical of high energy environments with loose sandy, well sorted to slightly muddy substrates in intertidal to shallow subtidal zones. The other trace fossils represent the Cruziana ichnofacies, which is typical of subtidal, poorly sorted and soft substrates, from moderate energy to low energy environments between the fairweather and storm wave base.The Kocayayla Group was deposited at an early stage in a shallow marine stable shelf condition. The shelf subsided in a later stage and was affected by normal faults along which mafic and felsic volcanic rocks erupted. The volcanic activity had ceased and a shallow marine clastic sedimentation took place in the final stage of the shelf development. The Kocayayla Group was deformed and metamorphosed before the deposition of the trilobite-bearing Middle-Upper Cambrian succession.     

北京昌平青白口系痕迹化石

北京昌平青白口系长龙山组和景儿峪组发育有丰富的痕迹化石,其中长龙山组产有Ｈｅｌｍｉｎｔｈｏｐｓｉｓ ｓｐ．,Ｃｏｃｈｌｉｃｈｎｕｓ ｓｐ．,代表潮下低能环境;景儿峪组产有Ｓｋｏｌｉｔｈｏｓ ｓｐ．和Ｍｏｎｏｃｒａｔｅｒｉｏｎ ｓｐ．,代表高能潮间环境。