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Trace fossil terms and definitions used in this paper.

Trace fossil terms and definitions used in this paper.

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Most studies of insect traces on fossil bone deal with one or two trace morphs found on isolated bone fragments, making it difficult to identify the trace-maker and its behavior. We report the discovery of a suite of insect traces on an articulated Camptosaurus dinosaur skeleton that permits the identification of the trace-maker and interpretations...

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Context 1
... facilitate description and discussion we list our trace terms and definitions (Table 2). Because many of the traces are part of a continuum, divisions between some of the morphs are artificial, but required for clarity. ...
Context 2
... pits have relatively smooth margins (Fig. 1B, lower right), whereas less deep pits contain fine grooves (Fig. 1 B-G). Shallow pits, pits less than 0.5 mm deep (Table 2), are comparatively uncommon, but their morphology ( Fig. 1F-G) is critical to understanding pit development. The rims of shallow pits are roughly the same shape and size as regular pits. ...
Context 3
... shallow pits consist of a small pit within a larger, shallow, flat-bottomed depression marked by radiating grooves (Fig. 1 F-G). Shallow bores, which are relatively uncommon (n = 25), have the same form as pits, except they are deeper and have parallel sides (Table 2). ...
Context 4
... cross section, bores (Table 2) are generally the same shape and size as pits and shallow bores, with the largest cross section being 6.5 × 4.5 mm. The bores (tunnels) enter bones mainly via articular surfaces ( Fig. 2 A, C, G, H-K), but some enter through thin skull elements or pass through ribs. ...
Context 5
... articular surfaces of nearly all bones of BYU 17945 are impacted by furrows with the destruction most apparent on large articular surfaces of appendicular elements ( Fig. 1 A, Fig. 2 A-C) or vertebrae (Fig. 2 D-K). Furrows (Table 2) are sinuous canals 1.8-8 mm wide that may remove a centimeter or more of bone ( Fig. 2 A-B). Broad pits are common along the bottoms of the furrows (Fig. 2 A, C, J), and some are contiguous with borings ( Fig. 2 C, center, and G, left margin). ...

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Citations

... They are direct evidence of interaction between insects and vertebrate remains and may contribute to the reconstruction of past environments, based on the identification of possible producers for the traces and the behaviors that generated them [ 1,2 ]. Insect bioerosion is relatively common in bones from Jurassic to Quaternary [2][3][4] ], but Triassic examples are considerably rare and up until now have only been reported for the Brazilian Santa Maria Supersequence (SMS) [5][6][7], making this unit a crucial landmark for the study of the origin and evolution of bone exploration by insects. Here, we report new observations of already analyzed specimens [ 8 ] and new examples of bioerosion traces in bones from the SMS and discuss their importance in the understanding of ethological diversity of bone exploration by insects in the early Late Triassic. ...
... Serrano-Brañas et al. 2018b, 2020. The small size of these traces is indicative of borings produced by invertebrates, of which only insects are known to damage bone in terrestrial environments (Britt et al. 2008). It is well known that bone modifications are different depending on the insect producer and its life stage; therefore, studies based on the damage succession of distinct insect groups present in a carcass can help elucidate decomposer-level diversity under local paleoclimatic conditions (Rogers Britt et al. 2008;Bader et al. 2009;Saneyoshi et al. 2011;Huchet et al. 2013;Xing et al. 2015;Parkinson 2016). ...
... The small size of these traces is indicative of borings produced by invertebrates, of which only insects are known to damage bone in terrestrial environments (Britt et al. 2008). It is well known that bone modifications are different depending on the insect producer and its life stage; therefore, studies based on the damage succession of distinct insect groups present in a carcass can help elucidate decomposer-level diversity under local paleoclimatic conditions (Rogers Britt et al. 2008;Bader et al. 2009;Saneyoshi et al. 2011;Huchet et al. 2013;Xing et al. 2015;Parkinson 2016). ...
Chapter
Reconstructions of deep time environments traditionally were based on the analyses and interpretations of fossil organisms in deep time. However, interpretations based exclusively on the morphologies and diversities of ancient organisms and placed within the framework of ecologically similar extant analogues can lead to errors related to time-averaging and spatial mixing rates. To develop a more robust paleoenvironmental analysis, modern paleontology has embraced taphonomic and sedimentological data in combination with organismal data to best understand temporal, spatial and evolutionary patterns in the fossil record. In this chapter, three case studies are presented regarding sedimentary-driven environmental interpretations carried out in the Upper Cretaceous Cerro del Pueblo Formation of the state of Coahuila, Mexico. Here, taphonomic and sedimentological data are interpreted together alongside the faunal and floral fossil associations, thus allowing the reconstruction of several terrestrial environments that developed in one of the southernmost regions of North America during the Campanian (~72.5 Ma).
... The middle (III-V) cervical vertebrae of YLSNHM 01003 are penetrated by two sinuous burrow casts from the right side ( central sutures. Each with the diameter of approximately 2 cm, the burrows appear to be a trace made by beetles, possibly foraging on soft tissues, laying eggs, or digging pupation chambers (Britt et al. 2008;Kirkland and Bader 2010;Xing et al. 2013). Unlike feeding traces attributed to termites in other dinosaur skeletons (Britt et al. 2008;Kirkland and Bader 2010;Backwell et al. 2012;Xing et al. 2013;Li et al. 2016;Serrano-Brañas et al. 2018), there is no other apparent evidence for bone-foraging activities such as pitting, boring, or trenching across the bone surfaces around the burrow casts of YLSNHM 01003. ...
... Each with the diameter of approximately 2 cm, the burrows appear to be a trace made by beetles, possibly foraging on soft tissues, laying eggs, or digging pupation chambers (Britt et al. 2008;Kirkland and Bader 2010;Xing et al. 2013). Unlike feeding traces attributed to termites in other dinosaur skeletons (Britt et al. 2008;Kirkland and Bader 2010;Backwell et al. 2012;Xing et al. 2013;Li et al. 2016;Serrano-Brañas et al. 2018), there is no other apparent evidence for bone-foraging activities such as pitting, boring, or trenching across the bone surfaces around the burrow casts of YLSNHM 01003. A block of four distal caudal vertebrae is associated with YLSNHM 01002 (Fig. 5A, B). ...
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... In most cases the grooves are randomly oriented and scattered over large areas of the bone surface. When densely concentrated, the grooves give the bone an etched appearance, similar to traces reported both from fossil (Kaiser 2000;Britt et al. 2008;Backwell et al. 2020) and modern (Backwell et al. 2012) bones (Fig. 6B, E). Subparallel grooves appear in at least three different clusters, two of them in UFRGS-PV-1581-T #17 (Fig. 6F). ...
... At least three subcircular concentrations of grooves were observed; they resemble the pits reported by Laudet and Antoine (2004: fig. 2B), Britt et al. (2008: fig. 1D), and Parkinson (2022: fig. ...
... Bioerosion traces made in bones deposited in continental settings are mainly attributed to insects, especially dermestid beetles (e.g., Kitching 1980;Rogers 1992;Martin and West 1995;Britt et al. 2008) and termites (e.g., Watson and Abbey 1986;Kaiser 2000;Fejfar and Kaiser 2005;Backwell et al. 2020). Trace morphologies as those present on UFRGS-PV-1581-T, such as isolated and clustered grooves, have been reproduced in experiments using both termites (Backwell et al. 2012) and dermestids (Parkinson 2022) and, consequently, are not a precise source of information of the identity of the tracemaker. ...
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New bioerosion traces produced by insects in bones are reported from the Hyperodapedon Assemblage Zone of the Santa Maria Supersequence (Carnian, Brazil). The bones are assigned to a single rhynchosaur Hyperodapedon mariensis individual and among the traces, the ichnogenera Osteocallis (Osteocallis mandibulus, Osteocallis infestans, and Osteocallis isp.) and Amphifaoichnus (Amphifaoichnus isp.) are recognized, along with two morphotypes of indiscrete traces: clusters of grooves and borings. All the traces are assigned to the action of insects exploring the rhynchosaur carcass. Osteocallis and associated clusters of grooves are interpreted as feeding traces, but whether they represent necrophagic or osteophagic behavior is still uncertain. The lack of direct evidence for the ethological interpretation of Amphifaoichnus precludes its sole correlation with osteophagy, and other possibilities, such as the construction of temporary domiciles related to feeding or sediment moisture, are discussed. The traces analyzed here indicate that the insects explored a buried carcass, challenging the automatic association of Osteocallis and prolonged subaerial exposure of bones, placing insects as relevant taphonomic agents that affect the preservation of vertebrate carcasses. Additionally, the first appearance record of Amphifaoichnus is expanded back more than 140 Ma, indicating that complex behaviors employed by insects in bone exploration were already established in the early Late Triassic, shortly after the oldest records of invertebrate bioerosion in bones on continental settings.
... Producer. Generally, Cubiculum ichnospecies have been previously documented and interpreted as dermestid beetle borings in many palaeontological studies (e.g., Martin and West, 1995;Hasiotis et al., 1999;Laudet and Antoine, 2004;Britt et al., 2008;Bader et al., 2009;Saneyoshi et al., 2011). ...
... Campanian of the Qarn Ganah, Kharga Oasis (this study). This ichnospecies has been recorded from the Middle-Upper Jurassic of South Africa (Britt et al., 2008;Xing et al., 2015), Upper Cretaceous of Brazil, Morocco (Ibrahim et al., 2014;Francischini et al., 2016) to Pleistocene of Brazil (West and Hasiotis, 2007;Dominato et al., 2009). . ...
... Producer. Asthenopodinae mayfly larvae (Thenius, 1988a(Thenius, , 1988bBritt et al., 2008;Ozeki et al., 2020). In particular, the Asthenopodinae mayfly larvae, a subfamily of Polymitarcyidae, have been assigned to be the producer of the present ichnospecies (Thenius, 1979(Thenius, , 1988aJalvo and Andrews, 2016;Genise, 2017). ...
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The uppermost part of the Campanian Quseir Formation of Kharga Oasis, Egypt, contains a concentration of turtle skeletal remains in a lagoon setting. They appear as three successive horizons (I‒III), alternated between the variegated shales and the glauconitic mudstones and conglomeratic layers within the Hindaw Member. However, bones recovered in horizon III present a higher preservation potential than the others. Therein, turtle remains are represented by mostly complete shells, partial shells and many scattered and weathered shell fragments. The studied turtles lived in small ponds and marshes and were deposited as autochthonous to parautochthonous relics. Their bones display significant bioerosion signatures on both carapace and plastron. The morphological analysis of the bioerosion structures preserved revealed 11 ichnospecies, belonging to nine ichnogenera. Eight of these ichnotaxa are recorded for the first time in turtle bones of Egypt. The recognised bioerosional structures appear as borings, shallow chambers, grooves, and punctures produced by ticks, beetles, polychaetae worms, fishes/crocodile, and gastropods. They were attributed to the ichnogenera Karethraichnus, Cubiculum, Osteocallis, Radulichnus, Osteichnus, Osedacoides, Sulculites, and Machichnus. In addition, some bite marks assigned to Nihilichnus also occur on a carapace peripheral. This grade of bioerosion was likely caused by relatively long exposure time before the final deposition or burial. In some cases, borings may be produced during the host’s lifetime. The new material considerably expands the stratigraphic and geographic distribution of this trace fossil assemblage and reveals that their producers may have been able to survive in other paleoenvironmental conditions.
... Producer. Generally, Cubiculum ichnospecies have been previously documented and interpreted as dermestid beetle borings in many palaeontological studies (e.g., Martin and West, 1995;Hasiotis et al., 1999;Laudet and Antoine, 2004;Britt et al., 2008;Bader et al., 2009;Saneyoshi et al., 2011). ...
... Campanian of the Qarn Ganah, Kharga Oasis (this study). This ichnospecies has been recorded from the Middle-Upper Jurassic of South Africa (Britt et al., 2008;Xing et al., 2015), Upper Cretaceous of Brazil, Morocco (Ibrahim et al., 2014;Francischini et al., 2016) to Pleistocene of Brazil (West and Hasiotis, 2007;Dominato et al., 2009). . ...
... Producer. Asthenopodinae mayfly larvae (Thenius, 1988a(Thenius, , 1988bBritt et al., 2008;Ozeki et al., 2020). In particular, the Asthenopodinae mayfly lar-vae, a subfamily of Polymitarcyidae, have been assigned to be the producer of the present ichnospecies (Thenius, 1979(Thenius, , 1988aJalvo and Andrews, 2016;Genise, 2017). ...
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El Hedeny, Sara Mohesn, Abdel-aziz Tantawy, Ahmed El-Sabbagh, Mohamed AbdelGawad, and Gebely Abu El-Kheir ABSTRACT The uppermost part of the Campanian Quseir Formation of Kharga Oasis, Egypt, contains a concentration of turtle skeletal remains in a lagoon setting. They appear as three successive horizons (I‒III), alternated between the variegated shales and the glauconitic mudstones and conglomeratic layers within the Hindaw Member. However, bones recovered in horizon III present a higher preservation potential than the others. Therein, turtle remains are represented by mostly complete shells, partial shells and many scattered and weathered shell fragments. The studied turtles lived in small ponds and marshes and were deposited as autochthonous to parautochthonous relics. Their bones display significant bioerosion signatures on both carapace and plastron. The morphological analysis of the bioerosion structures preserved revealed 11 ichno�species, belonging to nine ichnogenera. Eight of these ichnotaxa are recorded for the first time in turtle bones of Egypt. The recognised bioerosional structures appear as borings, shallow chambers, grooves, and punctures produced by ticks, beetles, poly�chaete worms, fishes/crocodile, and gastropods. They were attributed to the ichnogen�era Karethraichnus, Cubiculum, Osteocallis, Radulichnus, Osteichnus, Osedacoides, Sulculites, and Machichnus. In addition, some bite marks assigned to Nihilichnus also occur on a carapace peripheral. This grade of bioerosion was likely caused by rela�tively long exposure time before the final deposition or burial. In some cases, borings may be produced during the host’s lifetime. The new material considerably expands the stratigraphic and geographic distribution of this trace fossil assemblage and reveals that their producers may have been able to survive in other palaeoenvironmen�t
... Producer. Generally, Cubiculum ichnospecies have been previously documented and interpreted as dermestid beetle borings in many palaeontological studies (e.g., Martin and West, 1995;Hasiotis et al., 1999;Laudet and Antoine, 2004;Britt et al., 2008;Bader et al., 2009;Saneyoshi et al., 2011). ...
... Campanian of the Qarn Ganah, Kharga Oasis (this study). This ichnospecies has been recorded from the Middle-Upper Jurassic of South Africa (Britt et al., 2008;Xing et al., 2015), Upper Cretaceous of Brazil, Morocco (Ibrahim et al., 2014;Francischini et al., 2016) to Pleistocene of Brazil (West and Hasiotis, 2007;Dominato et al., 2009). . ...
... Producer. Asthenopodinae mayfly larvae (Thenius, 1988a(Thenius, , 1988bBritt et al., 2008;Ozeki et al., 2020). In particular, the Asthenopodinae mayfly lar-vae, a subfamily of Polymitarcyidae, have been assigned to be the producer of the present ichnospecies (Thenius, 1979(Thenius, , 1988aJalvo and Andrews, 2016;Genise, 2017). ...
Article
Full-text available
The uppermost part of the Campanian Quseir Formation of Kharga Oasis, Egypt, contains a concentration of turtle skeletal remains in a lagoon setting. They appear as three successive horizons (I‒III), alternated between the variegated shales and the glauconitic mudstones and conglomeratic layers within the Hindaw Member. However, bones recovered in horizon III present a higher preservation potential than the others. Therein, turtle remains are represented by mostly complete shells, partial shells and many scattered and weathered shell fragments. The studied turtles lived in small ponds and marshes and were deposited as autochthonous to parautochthonous relics. Their bones display significant bioerosion signatures on both carapace and plastron. The morphological analysis of the bioerosion structures preserved revealed 11 ichnospecies, belonging to nine ichnogenera. Eight of these ichnotaxa are recorded for the first time in turtle bones of Egypt. The recognised bioerosional structures appear as borings, shallow chambers, grooves, and punctures produced by ticks, beetles, polychaete worms, fishes/crocodile, and gastropods. They were attributed to the ichnogenera Karethraichnus, Cubiculum, Osteocallis, Radulichnus, Osteichnus, Osedacoides, Sulculites, and Machichnus. In addition, some bite marks assigned to Nihilichnus also occur on a carapace peripheral. This grade of bioerosion was likely caused by relatively long exposure time before the final deposition or burial. In some cases, borings may be produced during the host’s lifetime. The new material considerably expands the stratigraphic and geographic distribution of this trace fossil assemblage and reveals that their producers may have been able to survive in other palaeoenvironmental conditions
... Serrano-Brañas et al. (2018) figured channel-like bioerosion similar to our material, but their specimens are more sinuous, therefore indicating that these structures had been used as external mining/harvesting traces. Furrows are regarded as one of the bone modifications that have been produced by insects, similarly to other structures such as holes, pits, notches, grooves, scratches, chambers and tunnels (e.g., Roberts et al., 2007;Britt et al., 2008;Bader et al., 2009;Huchet et al., 2013;Pirrone et al., 2014b;Xing et al., 2016;Parkinson, 2016;Serrano-Brañas et al., 2018;Cruzado-Caballero et al., 2021). Fig. 4F Referred specimens. ...
... Discussion. In the literature, the chamber, cavity, or even hole borings within fossil bones are most commonly interpreted as pupation chambers of dermestid beetles (e.g., West and Hasiotis, 2007;Britt et al., 2008;Bader et al., 2009;H€ opner and Bertling, 2017;Serrano-Brañas et al., 2018;Augustin et al., 2019). However, the described structure does not show a high degree of similarity with the most approaching structures, which are generally attributed to the ichnogenus Cubiculum in overall shape and size; therefore, the morphology is herein treated as an indeterminate specimen. ...
... Identity of the trace makers 6.1.1. Insects as trace-makers on bones Ichnofossil-bearing bones preserved in terrestrial environments from several Mesozoic and Cenozoic localities are nearly always attributed to pupation and/or feeding by scavenging insects (Britt et al., 2008;Augustin et al., 2021). It is well known that the different bone modifications depend on the insect trace maker and its life stage and are generally expressed as furrows, grooves, holes, notches, scratches, pits, chambers, and tunnels (e.g., Roberts et al., 2007;Britt et al., 2008;Bader et al., 2009;Huchet et al., 2013;Pirrone et al., 2014b;Xing et al., 2016;Parkinson, 2016;Serrano-Brañas et al., 2018;Augustin et al., 2019;Cruzado-Caballero et al., 2021). ...
Article
Trace fossils, representing the ichnospecies Cubiculum ornatus, C. inornatus, Cubiculum isp., and Cuniculichnus variabilis as well as three indeterminate morphological categories (thin branching grooves A and B, furrows and circular chambers), are here recorded on vertebrate remains from the mid-Cretaceous (lower Cenomanian) of the ‘Continental Intercalary’ deposits in the Gara Samani locality (central Sahara of Algeria). Apart from thin branching grooves (A and B), which are thought to result from root etching, the bioerosion traces recorded on the bones comprise circular chambers and furrows, which represent pupation chambers of scavenging insects, most likely attributed to dermestid beetles. These trace fossils are identified and described for the first time from the Cretaceous of Algeria. They add new data to the trace-fossil assemblages previously reported from other Cretaceous successions of North Africa (e.g., the Kem Kem beds in Morocco). Sedimentological clues indicate a braided fluvial system environment. Taphonomic analysis suggests that the vertebrate remains were most likely subaerially exposed for a prolonged time prior to being transported and finally buried. Therefore, most of the studied borings are likely the result of dermestid beetle larvae feeding on dry bones on dry lands; other borings represent plant etchings produced after the bones were partially covered by, or completely buried, in sediments.
... Depressions that differ in shape from carnivore or rodent bite marks as described by White (1992) were classified as four types of pits, (e)-(h); these damage types may have been caused by insects or other small animals. Similar pits or holes have been assumed to have been made by termites or cutworms who passed through the bones to build their nests or gnawed on the bones (Britt et al., 2008;Fernandez and Andrews, 2016). Pit (e) is an oval-shaped depression without jagged striae at the bottom or around the depression. ...
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To evaluate burial customs in the Jomon period of Japan, we observed the taphonomic sig‍natures of human bones from a secondary collective burial and compared them with those from in‍dividual burials. We compared the compositions of identified bone parts, degree of weathering, and damage of the bone surfaces based on two neighboring shell-mound sites, Gongenbara and Horinouchi in Chiba Prefecture; a famous secondary collective burial had previously been discovered at the Gongenbara shell-mound. The degree of weathering differed between the collective and individual burials, and also differed depending on the horizontal and vertical location within the collective buri‍al. ‍These differences imply that a variable degree of weathering plausibly occurred at or after the secondary burial, but not at the primary burial. Regarding bone surface damage, specific types of dam‍age were more frequently observed on the limb bones in the collective burial than in the individual burials, although different types of damage were observed in the skull assemblage. These findings sug‍gest that different burial processes could cause the secondarily reburied bones to be exposed to the air for a longer duration of time or more easily be accessed by insects or other small animals, which might have produced variable taphonomic signatures.
... 36. Paik, 2000Roberts el al., 2007;Britt et al., 2008;Bader et al., 2009. 37. Nous proposons ici le néologisme de« termitoturbation ». , 1978 ;McBrearty, 1990 ;Balek, 2002 ;Fowler et al., 2004. ...
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Au cours de la fouille puis lors de l’étude bioanthropologique du site de Qedemt, il est apparût que les vestiges osseux humains étaient extrêmement fragmentés, incluant des lacunes importantes de nombreux éléments squelettiques. L’étude taphonomique de ces ossements a révélé que leur altération n’était pas corrélée à quelconque événement de nature physico-chimique (pH du sol, météorisation (weathering), compression du sédiment…) mais à des agents dits « biotiques » et plus spécifiquement des insectes. Un examen attentif des vestiges osseux en laboratoire à la recherche d’une « signature taphonomique » a permis de démontrer que les bioagents à l’origine de ces altérations ostéolytiques étaient des termites. La présence de nids subfossiles et actuels du Macroterminae Odontotermes rothschildianus (Sjöstedt) sur le site de Qedemt suggère que les nombreuses altérations osseuses répertoriées sur les restes humains sont majoritairement dues à ce taxon. L’étude entomologique réalisée sur le site a permis de révéler la présence d’une seconde espèce de termite, d’une sous-famille distincte, au sein de l’espace funéraire de Qedemt.