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Coprolites: a brief review with reference to specimens from the Rhaetic Bone Beds of England and South Wales
... The fundamental puzzle in the studies of coprolite is the difficulty in identifying the potential producer, which can be due to their nature and preservation. Also, that includes the methods used to deduce them with their producer, which were done by inferring with various forms of relationship based on stratigraphy and geographical relationships, as well as on neoichnology studies 7,23,54,55 . Such problems similarly arose in our context as well, and the materials were collected from a stratum that were interpreted as lake deposit margins, thus, suggesting an amphibious or aquatic producer. ...
... Above all, and more importantly, that the shape of the coprolite has to be intact in order to represent the shape of the internal intestine of the producer, whereby, anatomically it can lead to a certain biological aspect and digestive system of the organism. Despite these, there are on-going controversies on the origin of the spiral shaped bromalites in regards to whether or not they signify fossilized feces, or they are the cololite that was formed within the colon 6,21,23,61,62 . ...
... Most of the Tatal's coprolites are pink-whitish in color, which is highly associated with the presence of calcium through its carnivorous diets [93][94][95][96] . The dark colors can also be due to the presence of iron or it could also be due to complete phosphatisation 23,27 . However, a large part of the colorations was influenced by diagenesis 27,28 . ...
In this paper, seven coprolites from the Lower Cretaceous of Tsagan-Tsab formation have been described. Thus, producing a significant contribution to what we perceived as the first detailed study of coprolites from the Mesozoic deposits in Mongolia. The collected coprolites encompass a total of six spiral amphipolar and one scroll coprolites. We prominently identified four new coprolite ichnotaxa, such as: Hyronocoprus tsagantsabensis and Hyronocoprus hunti, to which both are ichnosp. nov.; followed by Megakalocoprus barremianensis and Scrollocoprus tatalensis, where both are ichnogen. et ichnosp. nov. Notably, CT scans revealed that all specimens showed various amounts of bony inclusions and scales, hence, deducing that the producers could have had a low acidic digestive track and were unable to dissolve bone matters. Moreover, SEM–EDS analysis concluded its carnivorous nature, thus, pointing towards piscivorous diet. The small sized Scrollocoprus is considered to be the second findings of Mesozoic era’s scroll coprolites, which contain possible plant pollens, a complete infraorbital bone, clusters of bone fragments and rhomboidal-shaped ganoid scales of the prey; and bioerosional scars have been observed on the surface. We suggest those amphipolar spiral ichnotaxa were produced by Asipenceriformes, with Pholidophoriformes as the prey, while Scrollocoprus represents fecal excrement of underived fish, possibly of sarcopterygian origins.
... In later studies, heteropolar edge and knot morphotypes were introduced 15 . With that, it was generally agreed upon that spiral coprolites are indeed producer of animals with a complex spiral valve intestine, such as the sharks, rays, lung shes and maybe ichthyosaurs 4,16,17,18,19 ; and are closely associated to aquatic environment; and generally, rapidly buried. ...
... The fundamental puzzle in the studies of coprolite is the di culty in identifying the potential producer, which can be due to their nature and preservation. Also, that involves the methods used to deduce them with their producer were done by inferring with various forms of relationship based on stratigraphy and geographical relationships, as well as on neoichnology studies 7,19,46,47 . Such problems similarly arose in our context as well, and the materials were collected from a stratum that is interpreted as lake deposit margins, thus suggesting an amphibious or aquatic producer. ...
... Among all, and more importantly, is that the shape of the coprolite has to be intact in order to represent the shape of the internal intestine of the producer, whereby, anatomically it can lead to a certain biological aspect and digestive system of the organism. Despite these, there are on-going controversies on the origin of the spiral shaped bromalites, on whether it signify fossilized feces or it was the cololite formed within the colon 6,17,19,53,54 . ...
Seven coprolites from the Lower Cretaceous of Tsagan-Tsab formation have been described, thus making this paper a significant contribution to the study of the firstly recorded coprolites from the Mesozoic deposits in Mongolia. The collected coprolites comprised a total of five morphotypes, whereby six of them are considered to be spiral amphipolar and one as rod shape. CT scans have also revealed that all specimens showed various amounts of bony inclusions and fish scales of unknown affinity, hence deducing that the producers may have had a low acid digestive track and were unable to dissolve bone matters. Moreover, SEM-EDS analysis concluded its carnivorous nature. In addition, all coprolites exhibited bony elements of identical morphological structures, which indicated that the producers might have preyed on the same type of organism. Specimen IVPP V 27545 is the only non-spiral coprolite and the smallest compared to the rest and contain plant pollens, an infraorbital bone and clusters of bones of the prey; bioerosional scars are observed on the surface. We suggest that the spiral coprolites were produced by Asipenceriformes with ichthyophagous diet, while the non-spiral rod like coprolite might possibly belong to smaller vertebrate organisms that can’t be identified by us in this study.
... Coprolites were mentioned in numerous papers listing the faunal components of the Rhaetian bone bed and Penarth Group sediments in general from a wide range of localities in subsequent literature. No serious attempt was made to characterise the Rhaetian coprofauna in any way until Duffin (1979), who focused on specimens from the conglomeratic bone bed at Aust Cliff. Duffin recognised four broad morphotypes and discussed the identities of potential producers. ...
... isolated from the surrounding sediment. This coprofauna can be used to update the classification and discussion given by Duffin (1979) and Swift and Duffin (1999), be subject to modern nondestructive imaging techniques and form the basis for simple statistical analysis with a view to elucidating more of the palaeoecological factors at play during the Late Triassic. ...
... The external surface of the coprolite possesses occasional longitudinal and transverse striae, but no spiral structure is visible. Some specimens have a "phosphatised skin", presumed to be a consequence of diagenesis (Duffin, 1979), and only visible in transverse sections of incomplete coprolites. Relatively homogenous overall, the coprolitic matrix varies from coarse-grained, resulting in an irregular external surface, to fine grained, giving rise to a smooth surface. ...
Coprolites from the Rhaetian bone beds in southwest England can be assigned to crustaceans and fishes. Here, we report crustacean microcoprolites, including Canalispalliatum and Favreina, the first records from the British Rhaetian, from Hampstead Farm Quarry near Bristol, evidence for diverse lobsters and their relatives not otherwise represented by body fossils. Further, we identify five fish coprolite morphotypes that differ in shape (cylindrical, flattened) and in presence or absence of a spiral internal structure. Many coprolites show bony inclusions on the surface, often relatively large in proportion to the coprolite; these show little or no evidence for acid damage, suggesting that the predators did not have the physiological adaptations of many modern predatory fishes and reptiles to dissolve bones. CT scanning has revealed the nature, packing and identity of inclusions within the coprolites, mainly fish scales, and some coprolites can contain more than twenty. An extraordinary discovery in one coprolite comprises a single sculptured skull element of the large bony fish Severnichthys together with two caudal vertebrae of the marine reptile Pachystropheus: did the coprolite producer, likely a fish, scavenge some flesh from the head of Severnichthys and then bite off the tail of the reptile? Assigning coprolites to producers is difficult, but it seems that Gyrolepis was preyed on by nearly every predator. The faunas and trophic relations revealed by the coprolites show that this was a modern-style marine ecosystem, with abundant crustaceans and several species of durophagous fishes, evidence for an early stage in the Mesozoic Marine Revolution.
... Coprolites are very common in the bone beds of the Westbury Formation in southwestern England, and many specimens are preserved in museum collections (e.g., Buckland 1835;Duffin 1979;Storrs 1994;Martill 1999;Swift and Duffin 1999;Gallois 2007;Hunt et al. 2012a, b: Fig. 12.12). There are more than half a dozen morphotypes in the Westbury coprofaunas, including both spiral (amphipolar and heteropolar) and nonspiral forms, Liassocopros hawkinsi, Saurocopros bucklandi and Strabelocoprus pollardi (Duffin 1979;Swift and Duffin 1999;Hunt et al. 2007Hunt et al. , b, c, 2013b fig. ...
... Coprolites are very common in the bone beds of the Westbury Formation in southwestern England, and many specimens are preserved in museum collections (e.g., Buckland 1835;Duffin 1979;Storrs 1994;Martill 1999;Swift and Duffin 1999;Gallois 2007;Hunt et al. 2012a, b: Fig. 12.12). There are more than half a dozen morphotypes in the Westbury coprofaunas, including both spiral (amphipolar and heteropolar) and nonspiral forms, Liassocopros hawkinsi, Saurocopros bucklandi and Strabelocoprus pollardi (Duffin 1979;Swift and Duffin 1999;Hunt et al. 2007Hunt et al. , b, c, 2013b fig. 12). ...
The diverse ichnofaunas of the Late Triassic have been studied for almost 200 years. During the Late Triassic, facies favorable for the preservation of trace fossils were the result of low sea levels, monsoonal climates and the development of extensive depositional basins as Pangea began to fragment. The most abundant vertebrate trace fossils in the Late Triassic are tetrapod tracks, including Brachychirotherium, Chirotherium, “Parachirotherium,” Synaptichnium, Atreipus, Grallator, Eubrontes, Banisterobates, Trisauropodiscus, Evazoum, Tetrasauropus, Pseudotetrasauropus, Eosauropus, Apatopus, Batrachopus, Rhynchosauroides, Gwyneddichnium, Procolophonichnium, Chelonipus, Brasilichnium and Dicynodontipus. There are five tetrapod footprint biochrons of Triassic age that can be identified across the Pangaean footprint record. Coprolites are the second most abundant vertebrate trace fossils in the Late Triassic and include Heteropolacopros, Alococoprus, Dicynodontocopros, Liassocoprus, Saurocoprus, Strabelocoprus, Malericoprus, Falcatocoprus and Revueltobromus. Coprolites are useful in biochronology in the Late Triassic. Consumulites, dentalites (new term for bite marks), and burrows are moderately common in the Late Triassic. Nests and gastroliths are rare. All groups of vertebrate trace fossils demonstrate different diversity and abundance patterns through the Phanerozoic. Most vertebrate trace fossils have their earliest occurrences in the Devonian. The early Permian is an acme for both tracks and coprolites. The Late Triassic yields abundant tracks and coprolites, and tracks are also common in the Early Jurassic. The Jurassic and Cretaceous represent the times with the greatest diversity of vertebrate traces (tracks, coprolites, consumulites, dentalites, nests and gastroliths). The Quaternary also represents a time of vertebrate ichnological diversity (tracks, coprolites, regurgitalites, nests and burrows).
... Triassic vertebrate coprolite records are known from many countries including but not limited to North America, South America, Greenland, Europe, Russia, Africa, China, Southeast Asia, Japan, Australia, and India (Duffin 1979;Northwood 2005, Hunt et al., 1998Hunt , 2018Wahl et al. 1998;Langer 2005;Hollocher et al. 2005;Burmeister et al. 2006;Klompmaker et al. 2010;Milàn et al. 2012Milàn et al. , 2021Bajdek et al. 2014, 2019Nakajima and Izumi 2014Brachaniec et al. 2015;Zatoń et al. 2015;Hansen et al. 2015;Niedźwiedzki et al. 2016;Luo et al. 2017;Qvarnström et al. 2019;Qvanstrom et al. 2021;Cueille et al. 2020;Hunt and Lucas 2021;Rakshit and Ray 2022). In India, the Upper Triassic vertebrate coprolite records are known from the Maleri Formation (Pranhita-Godavari Basin) and Tiki Formation (Son-Mahanadi Basin) of central India (Matley 1939;Sohn and Chatterjee 1979;Jain 1983;Vijaya et al. 2009;Rakshit et al. 2019;Umamaheswaran et al. 2019;Rakshit and Ray 2022;this article). ...
We record the first occurrence of a microcoprolite assemblage from the Upper Triassic Tiki Formation, Son –Mahanadi Basin, India. The microcoprolite assemblage has been assigned to four new ichnotaxa namely Cylindrocopros gondwanensis igen. et isp. nov., Elliptocopros rewaensis igen. et isp. nov., Convolucopros shadolensis igen. et isp. nov., and Cylindribulbocopros triassicus igen. et isp. nov. based on their external morphology, internal texture, and geochemical composition. Geochemical techniques confirm the phosphatic nature of the microcoprolites recorded from the Tiki Formation. Taken together, the associated fauna and the morphological comparisons between previously known coprolites from the Mesozoic-Cenozoic sedimentary sequences suggest that the Tiki microcoprolites were likely produced by carnivorous actinopterygians. The dentalites or biting traces observed on the external surface of the ichnotaxa recorded herein are the oldest records from the subcontinent. SEM analysis reveals the dominant presence of egg-like mineral spheres ‘microspherulites’, needle- and rod-shaped apatite crystals; however, the microcoprolites are devoid of biotic (floral and faunal) inclusions. Considering the abiotic crystals and geochemistry, we infer that the original faecal matter most likely underwent rapid precipitation during the early phase of mineralisation and had a very high potential for fossilisation.
... The methods employed to infer their origin are complex and encompass various approaches, including stratigraphy, geographical relationships, chemical analysis, and neocoprology studies. 28,29,57,[61][62][63] Coprolites often contain valuable dietary residues, making them akin to Lagerstätte, which are frequently underrepresented in the fossil record. It is widely acknowledged that certain coprolites serve as exceptional preservation sites, constituting Konservat-Lagerstä tten. ...
This study examines 55 coprolites from the Na Duong Basin to reconstruct the paleoenvironment. Coproecology sheds light on understanding the complex prey-predator relationships, trophic dynamics, and ecosystem evolution. Through quantitative and multidisciplinary analysis, the putative coprolites were attributed to crocodilian producers, leading to the establishment of a new ichnogenus and species, Crococopros naduongensis igen. et isp. nov., based on distinct characteristics and comparisons. The study provides compelling evidence of an ancient river or lake-like environment dominated by diverse crocodilian fauna, indicating a thriving food chain in the Na Duong Basin. The findings also highlight the remarkable richness of ichnofauna, fauna, flora, and the presence of a favorable climate, confirming the area as a significant fossil Lagerstätte in Southeast Asia. Overall, this study offers a unique snapshot of the past, providing valuable insights into the regional ecosystem and significantly contributing to our understanding of paleoenvironmental conditions and biotic interactions.
... The most di cult part of a coprolite study is to infer the possible producer, and we deem that as challenging due to their nature and preservation conditions. The methods used to deduce their producer are also quite complex, which are done by inferring with various forms of possible links that are based on stratigraphy and geographical relationships, examining their chemical constituents, and as well as on neocoprology studies 28, 29,54,51,55,56 . It is also known that much of the dietary residues are often embedded within the coprolite; hence, some authors have regarded coprolites as Lagerstätte within, which have often been under-represented in fossil records. ...
... The most di cult part of a coprolite study is to infer the possible producer, and we deem that as challenging due to their nature and preservation conditions. The methods used to deduce their producer are also quite complex, which are done by inferring with various forms of possible links that are based on stratigraphy and geographical relationships, examining their chemical constituents, and as well as on neocoprology studies 28, 29,54,51,55,56 . It is also known that much of the dietary residues are often embedded within the coprolite; hence, some authors have regarded coprolites as Lagerstätte within, which have often been under-represented in fossil records. ...
Reconstructing the paleoenvironment of Na Duong Basin by using coproecology is significant in understanding the intensive prey-predator relationships, the tropic relationship and the ecosystem evolution within the paleo-locality. A total of 55 measurable coprolites (IVPP V 27941/1 till IVPP V 27491/55) and numerous coprolite fragments from Na Duong fossil site were examined in this study. The putative coprolites were assigned to a crocodilian producer. Ichnosystematic studies further erected a new ichnogenus and species, which is Crococopros naduongensis ichnogen. et ichnosp. nov. for IVPP V 27491/46, based on a set of derived characters. Apart from quantitative analyses, a multi-disciplinary approach was also utilised for biogeochemistry analyses. The study has provided a rare and unique snapshot into the past where we believe an ancient river or lake-like environment has most likely existed in Na Duong Basin and was dominated by crocodilian fauna. We deemed that the Na Duong food chain was indeed ideal and healthy for the survival of the crocodilian species during that particular period of time with sufficient food sources. In addition, the study showed tangible evidence of the richness of ichnofauna, fauna, flora, the suitable climate, and paleoenvironment that supported Na Duong Basin as a fossil-Lagerstätte of Southeast Asia.
Abstract: Multiple spiral and non-spiral cylindrical coprolites
with profuse bone inclusions were recovered from a
metoposaurid-dominated bonebed in the Upper Triassic Tiki
Formation of India. The spiral coprolites are assigned to two
new ichnotaxa based on amphipolar and heteropolar micropolar
coilings. The non-spiral coprolites are internally devoid of
any spiral structure, mostly cylindrical, except for a few, which
are ellipsoidal. All of these coprolites contain randomly scattered
inclusions of partly digested and undigested skeletal fragments.
Based on external and internal morphology, and the
type of inclusions, the probable producers of these coprolites
are considered to have been carnivorous and/or piscivorous,
and included different types of chondrichthyans, dipnoans
and actinopterygians, as well as various semiaquatic and terrestrial
tetrapods. A web-like interconnectedness is evident,
which highlights dynamic and highly diverse animal communities
inhabiting the terrestrial, semiaquatic and aquatic realms
and dissolution of the inter-realm boundaries in the Upper
Triassic Tiki palaeoecosystem.
Key words: coprolite, India, Late Triassic, palaeocommunity,
vertebrate.
Multiple, small, cylindrical scroll coprolites having rounded and tapering ends and pertaining to a new ichnotaxon have been recovered from the Upper Triassic Tiki Formation of India. This is the first record of scroll coprolites from the Mesozoic. Based on cross‐sectional geometry, external surface textures, and internal morphology, these coprolites are subdivided into three morphotypes. The coprolites contain several kinds of undigested food material in the form of ganoid fish scales, teeth, lower jaw and skeletal remains of various osteichthyans, chondrichthyans, archosauriforms and indeterminate reptiles. These inclusions are embedded in the groundmass separated by thin mucosal layers. The groundmass contains abundant gas vesicles, and secondarily‐filled shrinkage cracks. EDS analysis shows that the overall composition of the coprolites reflects Ca, P, C and O, suggesting calcium phosphate mineralogy, though other elements such as Fe, Mn, Al, Si are present in lesser proportions. Based on their similarity with the scrolled faeces of extant euryhaline hammerhead sharks, it is deduced that these coprolites were produced by euryhaline hybodontid sharks. At least two hybodontid taxa, Lonchidion and Pristrisodus, show high prevalence in the Tiki vertebrate fauna, suggesting that these were the possible producers. As the coprolite inclusions contain remains of other aquatic animals, these carnivorous hybodonts constituted the dominant predators of the Tiki aquatic ecosystem.
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