Furthermore, coprolites did play an important role in preserving the traits of behavior 28 and it has been acknowledged that coprolites can provide salient analytical diagnostic on the feeding habits and dietary, prey-predator interactions, digestive physiology, diversity of the biota and environment, in which the organism lived to a certain extent on bacterial residues and DNA fragments 29 , 30 , 31 , 32 , 33 , This paper describes the biogenic structures, which herein attributed mostly to amphipolar spiral coprolites, found in the Lower Cretaceous of Tsagan-Tsab formation, western Mongolia.
Also, it is considerably the first detailed study of coprolite from Mongolia. At that time, they believed that the coprolites could have been produced by Choristoderans. A detailed descriptions of the specimens, followed by the succeeding comparison of the anatomical of the intestine features on related extant fishes based on past literatures have narrowed down the potential producer of the amphipolar coprolites, which could potentially belong to Asipenceriformes sturgeon and paddlefish see Discussion and interpretation.
Map showing the location of Tatal, in Mongolia. The coprolites were retrieved from the Tsagan-Tsab formation. The exact age for this formation is controversial. According to existing literature 37 , 38 , 39 , 40 , 41 , 42 , 43 , 44 , the Tsagan-Tsab formation could have range from the late Jurassic to Early Cretaceous, while Graham et al. Meanwhile, Krassilov 46 reported a Valanginian to Barrremian age by using plant fossils.
It has also been noted in Hasegawa et al. The Tsagan-Tsab formation is almost m in thickness, to which it is divided into upper and lower part, and consist of basal conglomerate to trough cross-bedded, coarse- to medium-grained sandstone, reddish or greenish shale, and calcretes 41 , 47 , It mainly consists of alluvial fan to lacustrine deposits 47 and forms a large shallow lake Over the years, Tsagan-Tsab formation has yielded numerous fossil fauna and among some, but not limited, include insects, mollusca, ostracods, fish, lizards, pterosaurs and indeterminate psittacosaurid, sauropods and theropods 36 , 42 , 44 , 49 , The specimens were excavated at Tatal, western Mongolia Fig.
Description: Hyronocoprus tsagantsabensis was identified through an incomplete holotype and two incomplete referred specimens. These three specimens Fig. However, with further examinations, we noticed the occurrence of disentanglement at the posterior end, which points to the fact that the actual length of the specimen is unknown.
In addition, the ends of all specimens are slightly tapered. Discussion: H. The exposed edge in the referred specimen IVPP V is not recognised as a flap edge, but rather a rough broken edge of an amphipolar coprolite. All specimens are considered smooth with inclusions.
Coprolites from Tatal. Scale bar equals 2 cm. Light microscopy photos of various surface adhesion on the coprolites. Scale bars as indicated. Etymology: Dedicated to Adrian P. Hunt, who named the genus in , followed with his major contribution in vertebrate coprolite studies by developing crucial and comprehensive notions and terms.
Diagnosis: H. It is generally four times longer than the holotype of H. It is The coprolite has six tight shallow coils that are not sharply separated. One end was slightly broken, but the other end has an acute spot.
The surface is smooth with visible cracks and borings. Also, some were seen to have six coils see 51 , Fig. We eliminated the fact that it could have belonged to a Mosasaur, due to its smooth surface texture and non-segmented lateral view. Thus, deducing the richness of biodiversity of Tsagan-Tsab fauna members, since they are substantially massive than any other spiral coprolite locality members.
The existent of a large spiral coprolite producer should be taken critically in order to understand the ecosystem of Tatal during lower Cretaceous. Thus far, H. Etymology: From the Latin mega big and from the Greek kalos rope , in reference to the appearance of an uncoiled rope and kopros dung. The spirals are spaced along the longitudinal section of the specimen and are not concentrated at one end. Megakalocoprus is similar to Kalocoprus see 53 , Fig.
In particular, there are more than four spirals in lateral view that lack tapering ends. It has been found that Megakalocoprus is three times larger than Kalacoprus. Discussion: The morphology of Megakalocoprus is almost identical to Kalacoprus and Hyronocoprus , as all three of them are distinguished by deep sulci spiral coils. They can be differentiated by the number of coils. Meanwhile, Megakalocoprus and Kalacoprus are discern by a comparison of their large size.
Due to their incompleteness, they have an antero-posterior length of more than The widest dimension measured at It also has one end that was flattened while the other hand has some damages, which does not affect the prediction of its total coils. There are at least 3 coils in a preserved state and does not extend above 4 coils. It has a rough surface with a length of Coils are clear but not sharply formed as the holotype specimen, which could be due to mucus covering the layer during defecation and quick burial.
The middle coil is slightly narrow than the end coil. One end is tapered, while the other end is flattened. Both specimens are similar in the number of coils. Notably, the largest specimen of Kalocoprus is Etymology: Scroll from the scroll-liked shape of the coprolite and kopros dung. Diagnosis: Small, anisopolar, cylindrical rod-like coprolite. It differs from the Tikicopros see 18 , Figs.
Possesses shallow linear or straight free edge that was almost worn out. Also, thin coil that is barely visible on the pointed tip can be seen. More significantly. Scrollocoprus differs from Eucoprus 7 , to which Eucoprus is perfectly cylindrical in shape, while Scrollocoprus has one end that is wider than the other.
Discussion: Scrollocoprus is the only scroll coprolite that was found from the Tsagan-Tsab Formation. Thus, making its discovery a noteworthy contribution towards understanding the fauna of Tatal during lower Cretaceous period. One side of the specimen has traces of bioerosional scars. Arrows indicates to the furrows. Blue line indicates the area where the cross section was made. Structure in yellow indicate the infraorbital bone.
I to L indicates the infraorbital bone in different angle. M and N shows the sensory canal of the infraorbital bone. Box in green shows true burrow while the rest are pseudo-borrows. Inset shows the position of the borings on the coprolite. Scales as indicated. It is a complete coprolite with a rounded cross section. One end is slightly tapered in a rounded shape, as compared to the other end that is tipped. Visible clusters of bone fragments and fish scales can be seen on its surface.
Discussion: At present, this ichnospecies is specifically known to be from the Tsagan-Tsab Formation of western Mongolia. As from the measurements, all collected coprolites vary in sizes Table 1. All specimens contained some degree of bone fragments and rhomboidal-shaped ganoid scales adhered to the coprolite surfaces Fig. Additionally, all specimens have smooth surfaces with little abrasion. Bite marks have also been found on specimen IVPP V 27,, in which these traces were short, parallel, shallow and isolated.
They have been formed from 3 furrows of roughly 3. Through CT scans and surface observation, we noticed that all specimens contained bone fragments and scales of varying degrees Fig. On the contrary, for specimen IVPP V 27,, we noticed a rather complete bone structure, such as the ribs and a segment of an infraorbital Fig.
All 4 samples showed high peaks of calcium and phosphorus. We have also described a potential pollen structure under SEM image Fig. No signs of abrasion were found on all of the coprolites. Coloration of the coprolites varied, thus, indicating they were buried in different sedimentary conditions. Meanwhile, specimen IVPP V 27, showed shallow coil deepness, therefore, this indicates that it was buried rapidly after excretion.
There are several pivotal evidences that corroborate to fecal origins of the Tsagan-Tsab Formation material: 1 basic morphology; 2 general shape and size 3 inclusions of the fecal matter; 4 high calcium and phosphorus content; 5 bioerosional scars; 6 borings and cavities; 7 concentric cracks. 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 , 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 , Spiral coprolites are producer of an animal with spiral intestine valves to increase the surface area of absorption, to slow down food movement in the bowel to maximise nutrient absorption, which has a significant strategy in surviving uncertain and harsh environment conditions 28 , 63 , Referring to past literature, it is generally agreed upon that the spiral shape is the only distinctively coprolite morphology, whereby it has been regarded as a true coprolite and can be correctly associated to the source animal, such as a range of fishes in particular 6 , 22 , Many primitive bony fishes except those of teleosts , fresh water sharks elasmobranches , coelacanths, Saurichthys , sturgeons and lungfishes are known to have the spiral valve intestine 51 , 64 , 65 , Also, Price 67 suggested that the amphipolar form could have been derived from palaeoniscoids.
The spiral coprolites collected for this study are mainly amphipolar in shape and one in scroll. As we know, generally heteropolar spiral coprolite are produced by sharks, which have complex spiral valves Therefore, we can exclude those in the family of elasmobranches as the potential producers and this can also be supported by the non-marine geological settings of Tsagan-Tsab Formation. But it is also noteworthy to mention that in previous studies, some workers have conducted observations on sharks that were kept in tanks, and were not been able to find any spiral fecal pellets.
Also, modern day sharks are totally unrelated to the ancient Permian pleuracanth sharks 6. Despite these, evidence of spiral fecal pellet can still be observed in some of the present-day fishes, such as the African lungfish Protopterus annectans , the Australian lungfish Neoceratodus forsteri , the long-nosed gar Lepisosteus osseus and the spotted gar Lepisosteus oculatus 6 , 70 , 71 , As for scroll coprolites, it is generally known to be produced by animal with longitudinal valves valvular voluta , whereby the valves naturally rolls in upon itself , in a way that it maximises nutrient absorption 8 , 9 , 17 , Gilmore 17 in his work mentioned that this type of valve must be primitive than the transverse valve valvular spiralis , which could be a modification of the previous ones.
This form is especially known to sharks of carcharhiniforms 73 , and it is evident that it could have been associated with sarcopterygian 53 , as well as anaspid and thelodont agnathans In this study, we recognised four new ichnotaxa for all the seven coprolite specimens. Assigning four new ichnotaxa does not conclude that the coprofauna are of four different types of animals.
Considering there are two distinct morphologies, which are the amphipolar spiral and scroll, we can deduce that at least two animals can produce these coprolites. But we have to carefully consider that diverse diets at different times for the same animal can often be variable, and soft fecal materials can range disparately after defecation, as well as taphonomy influence 74 , Specimen IVPP V 27, is remarkably huge and its producer should be a massive animal since large animals could produce small excrement, but small animals would not be able to produce big excrement 52 , Moreover, since there are no relevant fossils fauna found in the locality, we were unable to exactly identify the specific producer, rather, we deduced with relevant sources.
However, we do know that both amphipolar spiral and scroll coprolites can be attributed to certain types of fishes. As of these, we can conclude that the coprolites were produced by fishes in different sizes. Specimen IVPP V 27, differs from the rest by its shape and size, which makes prediction even harder, because it could be produced by either large or smaller animals. CT scans revealed that bony inclusions are evident in all of the coprolites Fig.
Specifically, bones in specimen IVPP V 27, are rather unaffected by the acidity of the digestive enzyme and these were evident by the presence of clusters of entire bones in the coprolite Fig. Furthermore, we identified an infraorbital bone of a fish.
CT scans revealed that the infraorbital bone has a sensory canal where it branches off at both ends Fig. With these, we can indicate that the producer of specimen IVPP V 27, poorly masticated the prey and also had a rather low gut digestion for food 28 , 55 , 76 , 77 , Specimen IVPP V 27, might belong to the first type of digestive strategy, whereby the producer has limited food processing in the mouth and the food stays in the digestive system for a short period of time.
This strategy is regarded to be efficient in conditions where food sources are abundant and the nourishment levels are sufficient The rest of the coprolites possibly belong to the second digestive strategy, as the bone content is fragmentary. This suggest the producer might have limited mastication with improved digestive assimilation and longer gut time to favour better absorptions of nutrients 55 , 80 , 81 , 82 , The third type of digestive strategy does not imply in our study.
It is also noteworthy to mention that the quantity of the inclusions is not correlated to the size of the coprolite, rather, it is dependable on the above-mentioned biological variables 28 , Carnivorous coprolites are normally composed of calcium phosphate and other organic matter, but it is important to be aware that the initial compositions are usually altered during fossilization processes Meanwhile, the excretion of herbivores is generally lacking in phosphates and their fossilization are mostly dependable of the mineral enrichment Through the morphological shape, the density of bone and scale inclusions on the surface from the CT scans, we can directly assume that these coprolites are inevitably produced by carnivorous organisms.
The reason for that was because the specimens were collected almost two decades ago and they were very well-kept in the archives throughout these years. As predicted, all 4 samples gave higher content of Ca and P, thus, there is no doubt that they are indeed fossilized fecal materials. At the same time, the SEM image potentially showed a pollen grain like structure.
Hollocher and Hollocher 86 documented a pollen image by using SEM, which brings our potential pollen image Fig. Although specimen IVPP V 27, is produced by an unidentified carnivorous vertebrate, it is common for carnivore coprolites to have plant remains within them. Also, it is known that spores and pollens are exceptionally well preserved within the encasement of calcium phosphate, which inhibits sporopollenin degradation Various reasons can be inferred for the presence of the pollen in specimen IVPP V 27,, to which it could either be by accident or by preying on an herbivorous animal.
Furthermore, it could also be through the adhesion on the excrement when the fecal is still fresh Pollens are in fact valuable information provider for paleoenvironment reconstruction, as well as for understanding the vegetation state of a particular era 87 , 89 , 90 , 91 , Hence, further palynology analyses are needed for future work.
EDS mineral composition and coprolite coloration can be correlated to a certain degree, in which it could also explain depositional origin The dark colors can also be due to the presence of iron or it could also be due to complete phosphatisation 23 , However, a large part of the colorations was influenced by diagenesis 27 , For example, on all of the burrow traces in specimen IVPP V 27,, only one traces showed burrowing holes, while the rest did not form a hole. While those specimens with internals, but without any traces on the outer surface, this can be explained by taphonomy processes, whereby the outer surface is covered with sedimentary and non-differentiable.
It was reported in Tapanila et al. In our study, we couldn't determine if the holes were constantly in diameter or not. Numerous tiny holes were visible on all of the coprolites surface, as well as within it, and these were most probably caused by gases within the fecal matters. Microvoids are quickly filled with water when fecal matter is excreted from the animal body, thus making the fecal becoming heavy and sinking to the lake floor Those lines only occurred once without any repetition on the rest of the surface.
On the other hand, deducing from the dented surface on the bitten marks, we predicted that the marks were most probably made by the biting pressures from the fish mandibles, which may indicate coprophagous behavior. The biting could have happened on the lake floor just before sedimentary deposition. Since the bitten marks are on the surface, this probably suggests unintentional scavenging and was eventually aborted during food search.
In general, coprolites can be transported from the original place through various modes 25 and this can be evident by the traces of abrasion 51 , Yet again, this supports our hypothesis that these coprolites were excrements in shallow waters, such as in the lake banks with little turbulence and current, where the fecal matter was dropped in-situ after excrement. As stated in previous literature , , radial and concentric cracks are also evident on the surface of specimen IVPP V 27,, therefore, these indicate that the coprolite was excreted on a very shallow environment where the water body was vastly evaporated and left for subaerial exposure before embedment.
This phenomenon caused the coprolite to dehydrate through the cracking, and shrinking occurred in a low magnitude process while retaining its overall shape 27 , 54 , Previous authors have also discussed that the cracks could possibly be due to synaeresis under certain conditions 27 , 54 , It has been frequently reported in records that almost all spiral coprolite fossilization from various Phenerozoic ages have occurred in low-energy shallow marine environments Feces that are being excreted in this humid environment have a higher chance of preservation due to the rapid burial, as well as on the acidity level of the water bodies 5 , 7 , , , There are also several crucial factors that are involved in fecal fossilization.
Among them, one of the most important criteria includes the content and composition of the fecal matter, and those of carnivorous diets tend to form coprolites than those who consumed an herbivorous diet As mentioned in Dentzien-Dias et al.
In accordance to this, we introduced the usage of phases to discuss the spiral coprolites morphologies in this study see material and methods. The phase concept of spiral coprolites disentanglement has been widely discussed in early days by various workers 6 , 22 , Although there are signs of disentanglement, we predict that the uncoiling on the surface was not by natural processes, but has been caused by a breakage after on.
Both of these two coprolites could have been large in actual size. Both of these specimens could have spent more time in water bodies before burial. Specimen IVPP V 27, does not provide any external information in regards of phases approach because of its non-spiral morphology. While it is also worthwhile to mention that none of them have spent sufficient time in the water bodies in order to possess the Phase 3 structure. Through these, we can also conclude that smaller coprolites are much complete while bigger coprolites tend to easily break-off.
Through the above morphological points, we predict that the amphipolar spiral coprolites could have belonged to groups of either prehistoric lungfishes or Acipenseriformes sturgeon and paddlefish. Another aim of this work is to portray the existence of possible prey-predation relationships from the collected coprolites. In order to narrow down the identity of the potential producer and possibly the prey, we looked into some related fauna list from past literature. Geological settings have indicated that the Lower Cretaceous Tsagan-Tsab formation is not only recorded in the area of Tatal, but also in other regions of Mongolia as well There are two possibilities on the deduced prey and predator, they are either of Asipenceriformes—Lycopteriformes relationship or Asipenceriformes—Pholidophoriformes relationship.
We suggest Pholidophoriformes as a much potential prey than the Lycopteriformes in the Tsagan-Tsab Formation, and the reasons will be explained thoroughly. It is also noteworthy to mention that the Tsagan-Tsab formations and the Yixian formation were similar in geological age.
In the same context, Jakolev 35 described Stichopterus popovi Asipenceriformes and recorded amphipolar spiral coprolites from the Aptian lacustrine of Gurvan-Eren Formation of Mongolia , a locality that is close to Tatal. Although there are differences in the geological period of Tsagan-Tsab and Gurvan-Eren Formation, it is highly possible that Asipenceriformes existed in these areas.
Furthermore, Asipenceriformes are shown to have spiral valves , and this can be further proven with the work of Capasso 64 on Peipiaosteus pani , thus, contributing to the morphology of the spiral coprolites. With these, we strongly suggest that the amphipolar spiral coprolites of Tsagan-Tsab Formation and for Gurvan-Eren Formation to belong to Asipenceriformes.
As for prey, we know from existing literature that there is a close relationship between Asipenceriformes and Lycoptera, as evident in the name Lycoptera-Peipiaosteus Fauna. Yondon et al. Another fact that was taken into account for the possible prey is the shape of the scales found in the inclusions, whereby Lycoptera are known for their cycloid shaped scales, while the ones in our specimens are more towards rhomboidal-shaped ganoid scales.
These facts crucially eliminate the possibilities of Lycoptera for the Tsagan-Tsab fauna. The size, shape of the scale and the nature of this fish fits well as a prey for the Stichopterus popovi Asipenceriformes. Through these interpretations, we can possibly infer that the spiral coprolites in our study might have belonged to Asipenceriformes and Pholidophoriformes as the prey, which could further affirm the occurrence of prey-predator inter-relationship in the Lower Cretaceous of Tsagan-Tsab Formation.
As for the sole scroll coprolite in this study, we do not intend to further deduce any detailed possibilities. Based on other works, chondricthyans origins or a sarcopterygian for scroll coprolites were suggested 18 , 53 ,but such deduction is difficult to be purported in our studies as there is a lack of such fossil materials in the locality and surrounding localities.
The chances of the underived producer to be a sarcopterygian is much higher than to be a chondricthyan, mainly due to its geological settings. The discovery of the single scroll coprolite can be a window opening to many paleontological questions for Tsagan-Tsab Formation. This study significantly contributes to the first detailed study of coprolites from the Mesozoic of Mongolia.
Specifically, we recognised four new coprolite ichnotaxa, such as: Hyronocoprus tsagantsabensis and Hyronocoprus hunti , to which both are ichnosp. Hyronocoprus tsagantsabensis , Hyronocoprus hunti and Megakalocoprus barremianensis are composed of amphipolar spiral coprolites, while Scrollocoprus tatalensis is scroll in shape.
Generally, through the SEM—EDS analyses and CT scans, we can conclude that all the studied coprolites have been produced by carnivorous organisms with piscivorous diet. Scrollocoprus tatalensis might be omnivorous consisting of animal and plant diets, as bony fish bones and pollen grain were evident, or it could have eaten preys with herbivorous diet.
All coprolites were in different sizes, inferring the producers were of different sized organisms. Additionally, the coloration, desiccation cracks, number of borings, cavities and coils deepness are different, indicating that these coprolites are buried under different taphonomy conditions. The producer of Hyronocoprus tsagantsabensis , Hyronocoprus hunti and Megakalocoprus barremianensis can be related to Asipenceriformes, while for Scrollocoprus tatalensis , we were unable to specifically link to any particular fish, but it could possibly be of sarcopterygian origins.
In addition, we predicted that the prey is from the Order of Pholidophoriformes. The study also shows that the ecology from where the coprolites were retrieved were once abundant in fish fauna. All of them came from the same locality, together with other fossil faunas, especially the pterosaurs The specimens are currently being housed at the Institute of Vertebrate Paleontology and Paleoanthropology, Beijing.
Five distinct morphotypes can be distinguished within the collected material. The term spiral coprolite is mainly divided into heteropolar, where the whorls or coils are concentrated at one end of the coprolite; while those of amphipolar, are recognized by the spiral which are spaced along the length of the specimen see 5 , 22 , Another category of coprolite in this study can be termed as rod-like or cylindrical elongated see 7 , Figs.
The term isopolar is referred to coprolite specimens with ends that are identical while anisopolar are for ends that are different in shape The definition of total length and coil length follows McAllister The coprolites were measured using the aid of a vernier caliper to the nearest millimeter by eye. Coloration is based on Munsell soil color chart Measurements, weight and general characteristics of the 7 specimens are summarized in Table 1.
In this study, we adapted the coil loosening approach to discuss the period of the excrement in the water bodies from the time of excrement to burial. This biological aspect was noted by some workers in the past. In Dean 70 , Williams 6 and Jain 22 , they observed that these excrements, when deposited in water bodies, tends to uncoil like ribbon like by hourly.
In this paper, we noted this process as phases to describe the morphology of the spiral coprolites. With this, we propose 3 phases in order to explain the period of the uncoiling, as explained by the aforementioned authors. Those include: Phase 1—Early phase of deposition, where all coils remain intact; Phase 2—Several hours after deposition, which some coils start to disentangle; Phase 3—After 24 h, where most coils have already loosened.
We conducted the scanning electron microscopy SEM coupled with energy-dispersive X-ray spectroscopy EDS on two specimens where the samples were easily obtained. For this, a tiny piece of sample was required by breaking it from the coprolite tip.
The specimens were then attached to a stub and coated with gold. The least damaging approach was considered in this process, which explains the reason on why only two samples were considered, and not all seven. A non-destructive technique using Computed Tomography CT scanning was used in examining the content and borings in the coprolites as well the production of a 3D model Supplemental video. All data generated or analysed during this study are included in this published article and its Supplementary Information files.
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Over the years practiced, palaeocoprology has been proven to be a bond for interdisciplinary fields by gathering various data from coprolites to understand the dietary structure, physical condition, genesis evolution, and phylogenetic status of producers. Accompanying with information about the ecological process, culture, and social evolution at the time, coprolites can further mirror every nuance of what has happened to the producers and environment in general at the time and place.
Due to the continuous microbial activities and deposition of other substances around coprolites as well as the change in taphonomic conditions such as temperature, humidity, and oxygen concentration, the non-contamination downstream analysis is a great challenge Appelt et al. In addition, an important issue in archaeozoology is the use of extant animal materials, such as bones, teeth, and feces, as control samples to interpret the results from the fossil record.
The following problem is that excellent records of a large number of reference individuals from controlled populations are difficult to obtain, especially for wild and extinct animals. Comfortingly, interdisciplinary technologies and tools have been deployed in the research of palaeocoprology to solve problems related to diet, evolution, and palaeoecology. In this situation, future research efforts should focus on the scientific and normative collection, transportation, preservation of coprolite, and the establishment of standardized procedures to remove contamination in subsequent experiments as well as to make a clear assessment of their negative effects on palaeocoprology, and eliminate them as much as possible.
Another step is expected to expand collaboration between archaeozoology, biology, and ecology to build more reliable frames of references, and more standardized experimental designs and methods. With the increasing discovery of coprolite-related ancient key sites around the world, coprolite researches will have greater value in the era of multi-interdisciplinary collaboration.
HX and XBZ designed this project. All authors contributed critically to the drafts and gave final approval for publication. The funding bodies contributed nothing to the study design, data analyses, data interpretation, or manuscript preparation. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
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They were first described by William Buckland in Prior to this they were known as "fossil fir cones " and " bezoar stones. Coprolites, distinct from paleofaeces , are fossilized animal dung. Like other fossils, coprolites have had much of their original composition replaced by mineral deposits such as silicates and calcium carbonates. Paleofaeces, on the other hand, retain much of their original organic composition and can be reconstituted to determine their original chemical properties. The fossil hunter Mary Anning had noticed that "bezoar stones" were often found in the abdominal region of ichthyosaur skeletons found in the Lias formation at Lyme Regis.
She also noted that if such stones were broken open they often contained fossilized fish bones and scales as well as sometimes bones from smaller ichthyosaurs. It was these observations by Anning that led the geologist William Buckland to propose in that the stones were fossilized feces and named them Coprolites. Buckland also suspected that the spiral markings on the fossils indicated that ichthyosaurs had spiral ridges in their intestines similar to those of modern sharks, and that some of these coprolites were black with ink from swallowed belemnites.
By examining coprolites, paleontologists are able to find information about the diet of the animal if bones or other food remains are present , such as whether or not it was a herbivore, and the taphonomy of the coprolites, although the producer is rarely identified unambiguously, especially with more ancient examples. The recognition of coprolites is aided by their structural patterns, such as spiral or annular markings, by their content, such as undigested food fragments, and by associated fossil remains.
The smallest coprolites are often difficult to distinguish from inorganic pellets or from eggs. Most coprolites are composed chiefly of calcium phosphate , along with minor quantities of organic matter. By analyzing coprolites, it is possible to infer the diet of the animal which produced them.
Coprolites have been recorded in deposits ranging in age from the Cambrian period to recent times and are found worldwide. Some of them are useful as index fossils , such as Favreina from the Jurassic period of Haute-Savoie in France. Some marine deposits contain a high proportion of fecal remains. However, animal excrement is easily fragmented and destroyed, so usually has little chance of becoming fossilized. Realising their potential as a source of available phosphate once they had been treated with sulphuric acid, he patented an extraction process and set about finding new sources.
The major area of extraction occurred over the east of England, centred around Cambridgeshire and the Isle of Ely [ 10 ] [ 11 ] with its refining being carried out in Ipswich by the Fison Company. This article incorporates text from a publication now in the public domain : Unsigned In Chisholm, Hugh.
Cambridge University Press. Wikimedia Foundation. Words nearby coprolite coprocessor , coproduce , coproduct , coprolagnia , coprolalia , coprolite , coprolith , coprology , coproma , coprophagous , coprophagy. But we bet not everyone knows these eight synonyms for poop. Get the low-down on what goes on down low with these words. How to use coprolite in a sentence Feces, especially from carnivores, can readily form into coprolite s. Fossilized dung from a dinosaur ancestor yields a new beetle species Nikk Ogasa June 30, Science News.
The secret to T. Smith February 8, Ars Technica. Fossilized excrement. Analysis of the fossilized animal and plant remains within coprolites provides important information about the diet and environment of ancient biota.
Coprolites are the. A coprolite is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour rather than morphology. The name is derived from the Greek words κόπρος and λίθος. They were. A coprolite is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in.