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Vertebrate coprolite studies: summary and prospectus
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... Tracks and footprints are the main vertebrate ichnofossils in this kind of sediment, which assist paleontologists in mechanical analysis of vertebrate movement of animal social behaviors (Lockley 1991;Cotton, Cotton, and Hunt 1997). Other vertebrate behavior trace fossils, such as nesting trace fossils (Grellet-Tinner et al. 2006) or coprolites (Thulborn 1991;Hunt, Lucas, and Lockley 1998;Hunt et al. 2012;Mancuso et al. 2018), are also interesting ichnofossils for paleoecological investigations. ...
The Mukdadiya Formation (Late Miocene-Pliocene) consists of alternations of red to brown sandstone and claystone layers in the Zagros Mountains Belt, northeastern Iraq. Two track-sites preserving bird and mammal tracks were recorded from the base of the formation in the Chamchamal area of the Kurdistan region in northeast Iraq. Avian tracks are large footprints with an average length of 25 cm that belong to Avipeda filiportatis. Mammal footprints imprinted by terrestrial cetartiodactyls belong to Pecoripeda amalphaea and Bifidipes velox, with cervids the most likely track makers. An unfamiliar large trace fossil on the studied slab, has symmetrical to asymmetrical marks, kidney or number-8-shaped and strong wrinkles on the surface. We interpret trace as having been produced by cetartiodac-tyls wallowing on the soft sediment surface. To accommodate this class of behavioral trace fossils; thus, we introduce a new fossilized behavior class, named "Laspichnia", which includes a vertebrate mud-bathing imprint on a soft sediment surface.
Coprolites are a highly informative but still underutilized proxy for understanding past environments, palaeodiets, and ancient human health. Here we provide a critical review of the history and current state of research in human coprolite analysis encompassing, macroscopic, microscopic, and biomolecular approaches. We present new data from a number of key sites which demonstrates how new multiscalar, multiproxy approaches can provide unique insights. Coprolites should be routinely collected and examined during excavations and integrated with other archaeological and palaeoecological evidence. Future research needs to focus on better understanding of coprolite formation as well as pre and post depositional taphonomy, a goal which can be achieved through interdisciplinary collaboration between geoarchaeology and organic geochemistry.
Novel fossil discoveries have contributed to our understanding of the evolutionary appearance of parasitism in flatworms. Furthermore, genetic analyses with greater coverage have shifted our views on the co-evolution of parasitic flatworms and their hosts. The putative record of parasitic flatworms is consistent with extant host associations and so can be used to put constraints on the evolutionary origin of the parasites themselves. The future lies in new molecular clock analyses combined with additional discoveries of exceptionally preserved flatworms associated with hosts and coprolites. Besides direct evidence, the host fossil record and biogeography have the potential to constrain their evolutionary history, albeit with caution needed to avoid circularity, and a need for calibrations to be implemented in the most conservative way. This might result in imprecise, but accurate divergence estimates for the evolution of parasitic flatworms.
The first vertebrate coprolite from the Upper Maastrichtian chalk of Denmark is described. The coprolite is 31 mm long, with a maximum diameter of 11 mm. Computed Tomography scanning of the specimen shows / revealed the internal structure to be tightly coiled, suggesting that the coprolite originates from a small shark with an estimated total length not exceeding 1.2 meters. Coprolites are normally found in abundance in marine deposits worldwide, so the lack of recorded findings from the Danish chalk probably reflects a collecting bias; collectors have simply failed to recognize them.
A significant number (more than 100) of brown to dark and silty, carbonate or pyrite-mineralized, in part organic carbon-rich, spherical or oval-shaped structures have been collected from the Upper Triassic (uppermost Norian-lower Rhaetian) sediments of the Lipie Śląskie clay-pit at Lisowice near Lubliniec town, Poland. Their geological context, morphology, content, geochemistry and association with skeletal remains suggest they are fecal masses of a sizable herbivorous tetrapod. The only large herbivore known from the site is a giant 5 meter-long dicynodont (Synapsida: Anomodontia), represented by numerous bones and also by large, oval-shaped footprints. The putative dicynodont coprolites were collected from mudstone and siltstone with numerous organic remains that were deposited in anoxic conditions. In addition, REEs and other trace element concentrations suggest that the burial environment and diagenesis of these coprolites were under anoxic conditions. SEM and thin section images of the coprolite matrix show numerous nests with pyrite (probably bacterial in origin) and large amount of mineral particles. The putative dicynodont coprolites contain also amorphous, dark organic matter, poorly preserved palynomorphs, small fragments of plant cuticle. Detailed characteristic of these coprolites reveals possible implications for the ecology and physiology of the source animal species. The δ 13C values of the gymnosperm cuticle and dark organic matter measured in three coprolites are − 23.4‰, − 21.2‰ and − 20.3‰, all average. The evidence from these coprolites suggests that dicynodonts processed plant soft elements into very small pieces, but wood fragments were found also in a mass accumulation in two coprolites.
Late Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) phosphatic coprolites from Central Europe are reviewed. These coprolites were predominately found in prey depot and commuting den hyena caves or at open air dens. In two cases pellets were found next to mammoth and straight-tusk elephant skeletons. Coprolites are identical in their shapes to those of modern African spotted hyenas (Crocuta crocuta crocuta (Erxleben, 1777). Feces consist of aggregates of pellets, built of several single droplets which are classified into seven different shapes and depend on their position within the aggregate. In most cases, coprolites contain small bone fragments of bone (compact or spongy). These bone remains cannot be attributed to the consumed prey without DNA testing. Pollen in coprolites do not allow a Pleistocene landscape reconstruction, because these originate from the intestines of the consumed prey and therefore reflect the prey's main food source. Radiocarbon dating of coprolites does not allow the determination of the exact time of extinction of the last hyenas of Europe. Fecal pellets are/were used by extant/extinct spotted hyenas to mark their dens and territories against other clans, lion prides and even wolf packs. A large number of coprolites and phosphatic layers, built of trampled excrements, are found at commuting or prey depot den sites (caves and open air), and are often important for distinguishing human and hyena den bone assemblages in Europe.
The new ethological class Digestichnia is introduced in the present article. It comprises all trace fossils
(and their recent counterparts) originating from the digestive process of animals, such as coprolites (feces),
regurgitalites (regurgitations) and gastroliths. The class is based upon a group within the unused classification
system for trace fossils proposed by Vialov in 1972.
Two fragmentary coprolites from the Lower Cretaceous Jydegaard Formation of Bornholm, Denmark, represent the first record of coprolites from continental Mesozoic deposits in Denmark. Both specimens contain fish scale bone fragments of unknown affinity. Based on morphology, inclusions and the potential producers found in the Jydegaard Formation, we suggest they were made by either a turtle or a theropod with a piscivorous diet. One specimen shows pits and grooves in the surface, as well as two deep cylindrical burrows, made by copropha-gous organisms.
A coprolite containing a tooth of an extinct shell-crushing shark, Ptychodus sp., was collected from the Blue Hill Shale Member of the Upper Cretaceous Carlile Shale in Ellis County, Kansas. The Ptychodus tooth lacks enameloid, probably as a result of etching through digestion in some animal. This specimen suggests that isolated ptychodontid teeth without enameloid which occur occasionally in other Upper Cretaceous deposits may be attributed to animal digestion.
Late Pleistocene fossils of the extinct Harrington's mountain goat, Oreamnos harringtoni, have been analyzed from eight localities in Grand Canyon National Park, Arizona. Unique finds of this species are the remains of hair, muscle and ligament, keratinous horn sheaths, and dung. Large, cuboid to sub-rectangular dung pellets are referable to the adult of the species. Pollen in dung indicates that the mountain goat frequented the caves during early to late spring and possibly portions of late winter and early summer. Plant material in the dung indicated that the major dietary components (33% to 47%) were grasses (Sporobolus, Festuca, Oryzopsis, and Agropyron); conifers (Picea and Pseudotsuga) also were important.
Screen-washing of unconsolidated fossiliferous sediments from the late Neogene Pipe Creek Sinkhole (Grant County, Indiana) yielded two coprolites. Maximum preserved diameter of both is about 26 mm, and both are apatitic in composition. Although one coprolite is largely amorphous internally, the other preserves remnants of hair and at least two teeth of a small carnivoran. The crowns of both teeth are highly corroded, and the enamel of one of the teeth has been completely removed. Although large turtles cannot be excluded as the scat-makers, the most likely candidate is a wolf-sized carnivoran, possibly a canid.
Coprolites are most common in the transition zone between the East Kirkton Limestone and the Little Cliff Shale. One hundred and twenty five samples are classified in five groups on the basis of size and shape. Within these groups distinctive subgroups are described. Many of the coprolites are believed to have been produced by fish, but some could have been produced by eurypterids and aquatic tetrapods.