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Hyena and ‘false’ sabre-toothed cat coprolites from the late Middle Miocene of south-eastern Austria
Historical Biology
Abstract
An association of eighteen coprolites (specimens 01–18) and one isolated coprolite (specimen 209,210) were found in a vertebrate fossil-rich paleosol at the Gratkorn site (south-eastern Austria; late Middle Miocene). The specimens consist mostly of calcium phosphate (apatite) and a matrix formed by microglobules. Coprolites 01–18 show cylindrical and spherical morphologies and are considerably smaller than the tube-shaped specimen 209,210, in which no inclusions were observed. In contrast, coprolites 01–18 contain numerous, highly altered bone fragments (sub-mm-sized long bones and several mm-sized trabecular bone remains) as well as hair imprints, plant detritus and palynomorphs. Based on composition, morphology, size, microstructure, and inclusions, and considering the body fossil record of this site, we assume the hyaenid Protictitherium and the barbourofelid Albanosmilus, as producers of coprolites 01–18 and 209,210, respectively. The preserved bone remains in specimens 01–18 suggest that Protictitherium fed on small vertebrates, but possibly also cracked bones of medium-sized animals. The hair imprints found were either from the hyaenid itself or its prey, while the plant material was probably ingested accidentally. The lack of inclusions in specimen 209,210 is related to the presumably hypercarnivorous diet of Albanosmilus, which was certainly the apex predator in this biome.
... isp. nov.; such longer morphotype was also found for the 'false' saber-tooth cat, Albanosmilus, from the late Middle Miocene of south eastern Austria [36]; similarly, Late Miocene coprolites from California, USA, studied by Wang et al. [34] and later ichno-systematically examined by Hunt and Lucas [35] led to the erection of a new ichnogenus and ichnospecies, Borocopros wangi, which shares size and morphology similarities with Cuocopros yuanmouensis igen. et. ...
... Additionally, scanning electron microscopy (SEM) images (Fig. 7) confirmed the presence of calcium phosphatic microglobule structures. Several researchers [26,27,36] have discussed the occurrence of these microglobules in their studies of carnivorous coprolites, proposing that they represent pseudomorphs formed after cocciform bacteria originating from the gastrointestinal microbiome of the defecating agent or the surrounding environment. These bacterial agents are considered crucial factors contributing to the enhanced preservation potential of the feces. ...
... These bacterial agents are considered crucial factors contributing to the enhanced preservation potential of the feces. However, it is important to note that the presence of calcium phosphatic microglobules in coprolites should not be solely relied upon as a diagnostic criterion for determining the potential dietary utility of the producer [36]. Hair inclusions are found in the Yuanmou coprolites and this may be of the defecating agent itself or from its prey. ...
This study reconstructs the Early Pleistocene paleoenvironment of the Yuanmou Basin through coproecology of the third member of the Yuanmou Formation. We examined 38 exceptionally well-preserved coprolites from a new fossil locality, and attributed the putative defecating agent to the hypercarnivorous diet canid, Sinocuon yuanmouensis through geochemical and quantitativeanalyses. A new ichnogenus and ichnospecies, Cuocopros yuanmouensis igen. et. isp. nov., was established based on distinctive characteristics. Multi-disciplinary analysis, including sediment palynology and lithostratigraphy, helped primarily reconstruct a significant climatic event during the early Pleistocene, coinciding with the emergence of Yuanmou Man during the fourth member of the Yuanmou Formation’s deposition. The findings provide insights into coexistence between canids, hyaenas, hominoids, and other fauna, revealing a rich paleoecosystem and food chain in the region’s history. This study contributes to understanding the complex ecological dynamics during this period in the Yuanmou Basin.
... isp. nov.; such longer morphotype was also found for the 'false' saber-tooth cat, Albanosmilus, from the late Middle Miocene of south eastern Austria [36]; similarly, Late Miocene coprolites from California, USA, studied by Wang et al. [34] and later ichno-systematically examined by Hunt and Lucas [35] led to the erection of a new ichnogenus and ichnospecies, Borocopros wangi, which shares size and morphology similarities with Cuocopros yuanmouensis igen. et. ...
... Additionally, scanning electron microscopy (SEM) images (Fig. 7) confirmed the presence of calcium phosphatic microglobule structures. Several researchers [26,27,36] have discussed the occurrence of these microglobules in their studies of carnivorous coprolites, proposing that they represent pseudomorphs formed after cocciform bacteria originating from the gastrointestinal microbiome of the defecating agent or the surrounding environment. These bacterial agents are considered crucial factors contributing to the enhanced preservation potential of the feces. ...
... These bacterial agents are considered crucial factors contributing to the enhanced preservation potential of the feces. However, it is important to note that the presence of calcium phosphatic microglobules in coprolites should not be solely relied upon as a diagnostic criterion for determining the potential dietary utility of the producer [36]. Hair inclusions are found in the Yuanmou coprolites and this may be of the defecating agent itself or from its prey. ...
This study reconstructs the Early Pleistocene paleoenvironment of Yuanmou Basin through coproecology of the third member of the Yuanmou Formation. We examined 38 exceptionally well-preserved coprolites from a new fossil locality, and attributed the putative defecating agent to the spectacular hypercarnivorous diet canid, Sinocuon yuanmouensis through geochemical and quantitative analyses. A new ichnogenus and ichnospecies, Cuocopros yuanmouensis igen. et. isp. nov., was established based on distinct characteristics. Multi-disciplinary analysis, including sediment palynology and lithostratigraphy, helped primarily reconstruct a significant climatic event during the early Pleistocene, coinciding with the emergence of Yuanmou Man during the fourth member of the Yuanmou Formation's deposition. The findings provide insights into coexistence between canids, hyaenas, hominoids, and other fauna, revealing a rich paleoecosystem and food chain in the region's history. This study contributes to understanding the complex ecological dynamics during this period in Yuanmou Basin.
Keywords: Coprolites, Early Pleistocene, Yuanmou, Coproecology, Cuocopros
... The putative tortoise (Testudinoidea) and snake (Serpentes) produces were assigned to the coprolites based on (1) the morphology compared to a large number of excrements of extant fish, amphibians, reptiles, birds and mammals and (2) the very limited number of inclusions. Gross et al. (2023) described 18 specimens from a vertebrate fossil-rich paleosol at the Gratkorn site (Serravallian/Sarmatian), Austria. The coprolites were assigned to hyaenid and barbourofelid producers based on the (1) morphology, size, (2) microstructure and inclusions and (3) the fossil record known from the site. ...
... The coprolites described by Peñalver and Gaudant (2010), Tomassini et al. (2019) and Gross et al. (2023) as well as one specimen by Antunes et al. (2006) are similar in age to the Pécs-Danitzpuszta specimens but were deposited in a different paleoenvironment. Collareta et al. (2022) and Godfrey et al. (2022) reported coprolites from a similar age and a locality with a similar fossil assemblage, including sharks, rays, cetaceans (Mysticeti and Odontoceti), seals, sirenians and crocodilians. ...
Thousands of coprolites have been collected from the Upper Miocene (Tortonian/Pannonian) sands of the Pécs-Danitzpuszta sand pit, one of the most important mixed Neogene vertebrate localities in Hungary. Although, the locality has been known for centuries, the coprolites have not been investigated in detail. We describe these fossils and explore their palaeoecological significance. Coprolites were examined with thin sections, X-ray powder diffraction and computed tomography. Their surface is mostly smooth, desiccation cracks were not observed. All coprolites mainly consist of apatite, indicating most probably a carnivorous diet. No evidence of herbivory has been found so far. The coprolites contain remains belonging to several groups: ostracod carapaces, bivalve shells, vertebrae from bony fishes, fish scales, as well as teeth. The CT scans show inhomogeneities in the matrix; however, in most cases they do not reliably indicate inclusions. The taphono-mical features suggest rapid burial and most likely an aquatic origin. The spiral morphotype could be attributed to fishes with spiral intestinal valves. Teeth of reef-associated fish taxa indicate Badenian age for some ARTICLE HISTORY
... Coprolites complement paleobotanical records as they also contain entomophile taxa (Djamali et al., 2011;Beck et al., 2019;Ochando et al., 2020). The study of hyena coprolites from caves has yielded very successful results (Gil-Romera et al., 2014;Gatta et al., 2016;Gross et al., 2023). However, guano from bats has attracted more attention of palynologists. ...
... This is explained by the fact that their faeces have a mainly phosphate composition (more than the average carnivore; Harrison, 2011) and that they deposit them in macro-latrines established inside caves (Pesquero et al., 2011), making their preservation more likely compared to other animals in the same climatic conditions (Kruuk, 1972;Larkin et al., 2000). The use of caves often means that they are also associated with human occupation, being present in very relevant archaeological sites such as Sierra de Atapuerca in Burgos (Pineda et al., 2017), Cova del Coll Verdaguer in Barcelona (Sanz et al., 2016), Calvero de la Higuera in Madrid (Arriaza et al., Several sites around the world and on the Iberian Peninsula have provided examples of coprolites from different producers found together in the same association (Abella et al., 2021;Brugal et al., 2012;Dentzien-Dias et al., 2018;Fiorelli et al., 2013;Gross et al., 2023;Sanz et al., 2016). Therefore, a detailed study of coprolites is necessary to identify the inhabitants of cave Quaternary environments even if there are no skeletal remains of them. ...
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Coprolites are very abundant in Quaternary deposits. However, they are often overlooked despite offering insights into both the surrounding environment and the species that produced them, particularly when skeletal remains are absent. We selected a Quaternary cave deposit from the Iberian Peninsula, Juan Labranz Cave, where an abundant and diverse coprolite collection was recovered, in order to develop a non-destructive study method that allows an accurate interpretation of the producer. Morphometric analyses of these specimens reveal two distinctive morphotypes associated with the two main carnivore occupations in the cave, hyaenas and bears. Morphotype I resembles hyaena coprolites in shape, texture, and bone inclusions, suggesting spotted hyaena as the most likely producer based on size. On the other hand, morphotype II deviates significantly, with a wrinkled texture and larger size, resembling bear faeces when eating carrion. Differences in bone fragment size and the presence of hair moulds further support these attributions. In conclusion, this study underlines the importance of considering taphonomy, morphology, size, and content in identifying coprolite producers, also high- lighting the relevance of these ichnofossils as indirect evidence of species and for the understanding of their ecological roles in Quaternary environments, where humans and other predators coexist.
The Eocene Pipestone Springs Main Pocket (Renova Formation, Jefferson County, Montana, United States of America) is a locality renowned for its diverse Chadronian (late Eocene; ∼38–33.9 million years ago) mammalian fauna and abundant coprolites. Two distinct coprolite size classes were previously identified in the trace fossil assemblage from which we selected representatives to investigate feeding behaviors and dietary selection of the producers. A subset of the selected coprolites was analyzed based on their compositional and taphonomic attributes using non-destructive x-ray tomographic microscopy in combination with more traditional methods including thin-section petrography, scanning electron microscopy, and energy dispersive spectroscopy. Among the features extracted in the tomographic data were skeletal fragments, including those showing evidence of bone-crushing; delicate hair molds; encrusted lithic fragments; and several irregular pores and cracks throughout the coprolites. Segmentation and volumetric renders permit quantitative assessment of the relative proportions of inclusions, revealing porosity as a primary volumetric element aside from the matrix and bone inclusions. There was no significant difference in the total volume of bone extracted between coprolite size class, though the smaller coprolites preserved a relatively higher volumetric proportion of undigested skeletal material. This multi-visualization approach provides a means to observe and evaluate differences in the coprolite gross morphology and inclusions across the two size classes, thereby offering valuable insights into the broader paleoecology of the Pipestone Springs Main Pocket coprolite producers and holding promise for comparable paleo-dietary studies of other coprolite-rich deposits.
The Early Pleistocene archeological site of Fuente Nueva-3 (FN3) preserves some of the oldest evidence of hominin presence in Western Europe, including a huge assemblage of Oldowan tools and evidence of butchering and marrow processing of large mammal bones. Moreover, there is also evidence of the regular presence of carnivores at the site, including a small proportion of bones that show tooth marks, the majority of which can be attributed to the giant, short-faced hyena Pachycrocuta brevirostris, and there are 220 coprolites, most of them from the Upper Archeological Level. In order to identify the defecating agent, we analyze here the coprolites and compare them with other specimens from the literature and with scats from zoo spotted hyenas (Crocuta crocuta). The morphology, color, size, and chemical composition of the FN3 coprolites allow us to attribute them to the hyena P. brevirostris, which is also represented at the site by fossil specimens. In addition, we evaluate the origin of the accumulation of coprolites and discuss on the role played by the scavenging hyenas in the accumulation and modification of the bone remains unearthed at the site, which allows evaluating the contribution of the giant hyena to this Early Pleistocene site. Finally, based on the lithology of layer 5 of the Upper Archeological Level, fine sands and clays deposited in a salt-lake environment, we hypothesize that this layer may have acted as a quicksand where large-sized animals like elephants were trapped and their carcasses lured scavenging carnivores.
The important vertebrate site Schönweg-"Brüchl" (SW St. Andrä im Lavanttal, Miocene, Lower Badenian, ~ 16 million years) yields a relatively diverse carnivore fauna with five species. While the dog bear Amphicyon cf. major and the marten-like Trocharion aff. albanense are already known in Austria, the viverride Forsythictis cf. aurelianensis, the small and basal hyena cf. Protictitherium sp. and the rare and special carnivore Sivanasua viverroides are new records for this country. This carnivore community lived in a humid, rather wooded environment.
A fragmentary mandible of the basal hyena cf. Protictitherium sp. from the Miocene of the Schönweg-"Brüchl" locality (Lavanttal, district Wolfsberg, Carinthia) still shows three milk teeth and the second molar in the process of eruption. Using modern X-ray computer tomography methods, the adult premolars p4, p3, p2, canine and probably i3-i2 can be then studied. The dental eruption sequence might be similar to that of Genetta genetta and other viverrids. If confirmed by new finds, the occurrence of the genus Protictitherium at Schönweg-"Brüchl" would be the first in Austria, but also one of the oldest in Europe.
The Middle Miocene (15.99–11.65 Ma) of Europe witnessed major climatic, environmental, and vegetational change, yet we are lacking detailed reconstructions of Middle Miocene temperature and precipitation patterns over Europe. Here, we use a high‐resolution (∼0.75°) isotope‐enabled general circulation model (ECHAM5‐wiso) with time‐specific boundary conditions to investigate changes in temperature, precipitation, and δ¹⁸O in precipitation (δ¹⁸Op). Experiments were designed with variable elevation configurations of the European Alps and different atmospheric CO2 levels to examine the influence of Alpine elevation and global climate forcing on regional climate and δ¹⁸Op patterns. Modeling results are in agreement with available paleobotanical temperature data and with low‐resolution Middle Miocene experiments of the Miocene Model Intercomparison Project (MioMIP1). However, simulated precipitation rates are 300–500 mm/yr lower in the Middle Miocene than for pre‐industrial times for central Europe. This result is consistent with precipitation estimates from herpetological fossil assemblages, but contradicts precipitation estimates from paleobotanical data. We attribute the Middle Miocene precipitation change in Europe to shifts in large‐scale pressure patterns in the North Atlantic and over Europe and associated changes in wind direction and humidity. We suggest that global climate forcing contributed to a maximum δ¹⁸Op change of ∼2‰ over high elevation (Alps) and ∼1‰ over low elevation regions. In contrast, we observe a maximum modeled δ¹⁸Op decrease of 8‰ across the Alpine orogen due to Alpine topography. However, the elevation‐δ¹⁸Op lapse rate shallows in the Middle Miocene, leading to a possible underestimation of paleotopography when using present‐day δ¹⁸Op—elevation relationships data for stable isotope paleoaltimetry studies.
The Gleisdorf Formation (Fm.) deposits in the clay pit at Gratkorn, Styria, Austria, are dated to 12.2–12 Ma, and are of late middle Miocene age (late Serravallian or Sarmatian). To reconstruct the paleovegetation and estimate the paleoclimate at this important vertebrate site, the palynoflora close to the boundary between the vertebrate-bearing layers of the Gratkorn Fm. and the overlying limnic clay deposits of the Gleisdorf Fm. was investigated. Using the single-grain method, 140 palynomorphs were identified. The palynoflora suggests that the paleovegetation was characterised by well-drained lowland and upland forests, riparian forest, and swamp forests. Depending on the dominating tree species, lowland and upland forests might have had closed or more open canopies. Open habitats included wet meadows and shrublands. In addition, conifers were present in the swampy lowlands and the forested uplands. The most prominent paleoclimatic signatures of the palynoflora indicate a fully humid warm temperate climate, with hot to warm summers and cool winters (Cfa-, Cfb-climate), and a seasonal climate with cool and drier winters and hot to warm and wetter summers (Cwa-, Cwb-climate). Our results align with existing studies bordering the Styrian Basin and support the presence of subtropical to warm-temperate vegetation around Gratkorn during the Sarmatian.
Pleistocene large carnivores from the Pampean Region have been studied almost always based on bone remains. Here we report the first coprolite of a large carnivore from the Late Pleistocene of the Pampean Region, Argentina. The coprolite was found associated with megafauna bone remains and it shows the typical shape and size of a carnivorous mammal, with a length of 240 mm and a maximum diameter of 39.59 mm. The coprolite contains two bones of the autopodium of an artiodactyl, and numerous dermal ossicles of giant terrestrial sloths. The coprolite's composition, size, and shape allow us to discard ursids, canids, and small felids. Ursid and canid feces contain abundant plant remains, which are absent in the coprolite described here. Feces of extant felids are smaller than the coprolite here as they do not exceed 130 mm in length and 30 mm in diameter. Based on size, shape, and bone inclusions, the discovered coprolite may be attributed to the Machairodontinae saber-toothed felid Smilodon. If correctly identified, this coprolite sheds light on predatory habits of Smilodon.
The present study deals with new material of carnivorans (Mustelidae, Mephitidae, Ailuridae, Potamotheriinae and Viverridae) from the basal Tortonian (Late Miocene, late Astaracian) hominid-bearing locality of Hammerschmiede (Bavaria, Germany). The small carnivoran fauna includes 20 species belonging to nine different subfamilies (Guloninae, Lutrinae, Mellivorinae, Potamotheriinae, Leptarctinae, Mephitinae, Simocyoninae, Genettinae and Viverrinae). The identified forms include: “Martes” sansaniensis, “Martes” cf. munki, “Martes” sp., Circamustela hartmanni n. sp., Laphyctis mustelinus, Guloninae indet., Eomellivora moralesi, Vishnuonyx neptuni, Paralutra jaegeri, Lartetictis cf. dubia, Trocharion albanense, Palaeomeles pachecoi, Proputorius sansaniensis, Proputorius pusillus, Alopecocyon goeriachensis, Simocyoninae indet., Potamotherium sp., Semigenetta sansaniensis, Semigenetta grandis and Viverrictis modica. The new species Circamustela hartmanni n. sp. is differentiated from the other members of the genus by its small size and the morphology of its dental cusps in the upper and lower carnassials. This is one of the highest reported taxonomic diversities for fossil small carnivorans in the Miocene of Europe, including also first and last occurrences for several genera and species. Additionally, the assemblage comprises some rare taxa such as Palaeomeles pachecoi and Eomellivora moralesi. An ecomorphological comparison of the discovered taxa reveals possible cases of competition and niche partitioning.
Excrement-shaped ferruginous masses have been recovered from the Miocene of
Turów mine in south-western Poland. These siderite masses have been the subject of
much controversy, having been interpreted either as being coprolites, cololithes or
pseudofossils created by mechanical deformation of plastic sediment. Here we
present the results of mineralogical, geochemical, petrographic and
microtomographical analyses. Our data indicate that these masses consist of siderite
and iron oxide rather than phosphate, and rarely contain recognizable food residues,
which may suggest abiotic origins of these structures. On the other hand, evidence in
support of a fecal origin include: (i) the presence of two distinct morphotypes
differing in size and shape, (ii) the presence of rare hair-like structures or coalified
inclusions and (iii) the presence of rare fine striations on the surface. Importantly,
comparative actualistic study of recent vertebrate feces shows overall resemblance of
the first morphotype (sausage-shaped with rare coalified debris) to excrements of
testudinoid turtles (Testudinoidea), whose shell fragment was found in the
investigated locality. The second morphotype (rounded to oval-shaped with hair-like
structures), in turn, is similar to the feces of some snakes (Serpentes), the remains of
which were noted in the Miocene of the neighborhood areas. Other potential
producers (such as lizards and crocodiles) and even abiotic origins cannot be fully
excluded but are less likely
Coprolites (mummified or fossilized feces), belonging to the group of ichnofossils, are fossilized remains of feces produced by animals. Various types of data from coprolites provide detailed evidence of the producer’s condition, like diet, intestinal microbiome, virus infection and parasites diseases. In addition, the palaeoenvironment information relevant to producers’ ecological niche can be drawn from taphonomy details the coprolites mirrored. At present, the phylogenetic clues of the producer’s population can be determined by advanced molecular biotechnologies. With the integration of multiple methods and techniques, coprolite has been widely accepted as an ideal material to study the diet, evolution, and palaeoenvironment of producers. In this paper, we reviewed the history of coprolite research, enumerated and interpreted the data recovered from coprolites, and explained their research value to palaeocoprology and evolutionary biology. Finally, we summarized the current directions of coprolite research and looked into its future prospects.
There is a long tradition of indirectly inferring feeding in fossil and Recent vertebrates from the functional morphology of bones and teeth. However, some trace fossils provide direct evidence of feeding, and the study of the ichnology of feeding dates to the 1820s–1830s. Trace fossils that document vertebrate consumption in all of its phases are dentalites (“tooth marks”), gastroliths and bromalites (includes coprolites, consumulites and regurgitalites, among others), and these trace fossils are key to understanding the evolution of nutrient acquisition and food processing in the evolutionary history of vertebrates. We review the fossil record and significance of these ichnofossils as they relate to feeding, which is to say we review the ichnology of vertebrate consumption. Beyond a review, we also indicate areas for further research, which are many, on dentalites, gastroliths and bromalites. Finally, we analyze our knowledge of the history of these trace fossils to identify critical biotic events and turning points in the evolutionary history of vertebrate consumption.
There is an extensive fossil record of vertebrate dentalites (“tooth marks”), principally on bony substrates, but also on invertebrate hard parts, coprolites, plants, lithic substrates and others, reviewed here for the first time. Two centuries of studies of dentalites show strong biases towards archeology, and towards dinosaurs as a result of the Taxophile Effect. This record merits more synthetic study in an ichnotaxonomic framework and the development of criteria for establishing inferences about behavior. Indeed, dentalites are of diverse paleoethological significance, including: (1) predation, including hunting strategies; (2) bite method and force; (3) dietary selection; (4) feeding; (5) scavenging strategies; (6) methodologies of bone accumulation; (7) trophic patterns; (8) intraspecific (agonistic) interactions; (9) tooth sharpening; and (10) bone and rock utilization for other purposes, including mineral extraction. What is now needed is a dentalite ichnology beginning with diverse documentation of the dentalite ichnofossil record, compilation and synthesis of the entire record, rigorous ichnotaxonomy and determination of analytical criteria for establishing inferences about the behaviors archived by tooth-mark ichnofossils.
Bromalites include regurgitalites, consumulites, coprolites, pabulites and digestilites. Regurgitalites are the least studied bromalites, the most difficult to identify, and their fossil record is strongly controlled by taxonomic and taphonomic factors. Bromalite pellets can represent coprolites or regurgitalites, and two-dimensional examples could be taphonomic artifacts (decayed specimens or physical concentrations). Many identified vertebrate regurgitalites were produced by fish or birds and are preserved in a limited range of environmental settings (e. g., aquatic low energy). Regurgitalites have diverse utility, including: (1) providing evidence of the evolution of predation and digestion; (2) analysis of taphonomy and sedimentary environments; (3) proxies for the presence of biotaxa; (4) loci for exceptional preservation; (5) biogeographic studies; (6) evaluating digestive processes of producers; and (7) evidence of the evolution of durophagy.
There are numerous descriptions of consumulites, reviewed for the first time here, but they are usually concealed within publications with a different focus. Consumulites give the most unambiguous dietary attributions of any bromalites and provide direct evidence of the nature of digestion and the structure of the digestive tract. However, the study of consumulites is in an early stage of development, though they also have great potential to provide direct evidence of aspects of patterns of digestion, such as: (1) assessing the chemistry of digestive systems by examining the etching and erosion of consumulite materials; (2) studying the evolution of the components of the digestive system; (3) analyzing dietary changes through ontogeny; (4) evaluating the evolution of diets within clades; and (5) identifying environmental tolerances. Consumulites may preserve a wide range of organic elements with a poor fossil record and thus can be Lagerstätten. In addition, consumulites can also preserve tissues of the gastrointestinal tract. The systematic study of consumulites will undoubtedly yield significant records of contained fossils, as has the recent focus on the contents of coprolites.
The term gastrolith refers to sand and/or gravel swallowed by an animal and retained in the digestive tract. Wings recently grouped objects with different origins under the single term gastrolith, so he proposed to add prefixes to the word gastrolith to create terms that identified their different origins: “bio-gastrolith” for the calcareous concretions formed in the bodies of some crustaceans; “patho-gastrolith” for concretions formed in the stomach pathologically; and “geo-gastroliths” for swallowed rock particles. These terms are unnecessary if gastrolith is restricted to the meaning we advocate, which is the same as Wing’s “geo-gastrolith.” Wings also advocated using the term
2
“exolith” for “stones” that might be gastroliths but that lack a convincing skeletal association. This term, however, enshrines the widespread misconception that highly polished “stones” are gastroliths, regardless of any skeletal association. It also lacks specificity, as any polished “stone” anywhere could be called an exolith. Thus, we do not use the term exolith. The distribution of gastroliths is very irregular in extant vertebrates and is often related to the presence of a muscular gizzard. Nevertheless, swallowing or not swallowing sand/gravel can be specific to some individuals within a species. A variety of functions have been suggested for gastroliths, only two of which are of significance: use in digestion to grind, pulverize and/or disintegrate food or use as ballast for buoyancy control. It seems likely that some crocodiles and marine mammals use/used gastroliths for buoyancy control, but all other gastrolith-bearing vertebrates appear to have used them in digestion. Identification of fossil gastroliths is only certain when the sand/gravel is found as a concentrated mass in an anatomically plausible position within the abdominal region of a fossil skeleton. There are various ways to polish stones, notably by the wind to make them ventifacts. Polished clasts identified as gastroliths are siliceous, mostly quartz or chert. These clasts could have been polished by wind and/or water, both before ingestion by an animal and/or after excretion or other removal from an animal’s digestive tract. Furthermore, how a stomach or gizzard would polish siliceous clasts (which are very hard) is also unclear, and such polishing does not occur in extant birds. Only a small minority of bona fide gastroliths, particularly those of plesiosaurs, are highly polished. There is thus an inability to establish when and where the clasts were polished, and inferring that they acquired their polish while gastroliths is not supported by actualistic studies of gastroliths. The idea that any highly polished clast is a gastrolith needs to be abandoned. Most records of gastroliths are from plesiosaurs, birds and some dinosaurs. Gastroliths are trace fossils in need of ichnotaxonomy that provide important insights into various behaviors. Gastroliths clearly are the work of an animal. Unlike eggs, for example, the sand/gravel that comprise gastroliths (our definition) is not made by animals–it is swallowed by, concentrated by, transported by and, in some cases, altered by animals. So, the trace-fossil status of gastroliths is unimpeachable. We advocate development of an ichnotaxonomy for gastroliths. Ichnotaxonomic names, however, should not be assigned to individual grains/clasts of sand/gravel of fossil gastroliths. We favor naming the entire gastrolith mass from a single abdominal cavity. Ichnotaxobases could be the number of gastroliths in the mass, their general petrographic composition, and their overall size, shape, surface texture and other shared morphological features. This likely would produce a workable ichnotaxonomy that recognizes ichnotaxa that are readily distinguished by morphological differences that are a direct reflection of varied behavior. Gastroliths provide important insights into various behaviors, notably diet, digestion, buoyancy control and habitat preferences.
After footprints, coprolites are the most studied vertebrate trace fossils, and they are the subject of an extensive and rapidly growing literature. Coprolites have an extensive fossil record that has proven potential to address a broad range of paleontological issues: (1) coprolites as proxy for biotaxa have utility in biochronology, biogeography and faunal turnover; (2) coprolites as trace fossils can delimit a hierarchy of ichnocoensoes and ichnofacies; (3) coprolites as end products of the gastrointestinal tract can provide evidence of digestive processes; and (4) internally, coprolites can be Lagerstätten that preserve a wide range of organisms with an otherwise poor fossil record.
Other bromalites are digestilites, pabulites and micturalites. Digestilites is a new term for materials derived from the digestive tract. They are subject to chemical and physical processes that result in characteristic damage that can be recognized in regurgitated or defecated material. Digestilites composed of invertebrate debris provide a particularly important insight into the evolution of durophagous fish and also constitute a significant sediment source, particularly in the Cenozoic. Many late Cenozoic (and some earlier) microvertebrate accumulations consist of digestilites, and this large topic deserves substantial study. Pabulites are fossilized food that never entered the digestive tract. Footprints, some nests and other traces also can record evidence of vertebrate predation and consumption. Mololite is a new term for tooth wear, which is a type of trace fossil.
There are taphonomic megabiases in the trace fossil record of vertebrate consumption. The bromalite and dentalite records are heavily skewed towards carnivores. There are also significant size-related biases, for example that small and large coprolites and regurgitalites are rare, as are small dentalites. Certain time periods demonstrate strong, geologically-based biases. For example, the continental flooding of the Late Cretaceous resulted in extensive trace (and body) fossils preserved in the Western Interior Basin of North America, not only in the sedimentary deposits of the seaways but also in the rocks that formed on the associated coastal plains. The first large sample of vertebrate ichnofossils related to feeding is from the Devonian, but the largest acme is in the Late Cretaceous, which resulted from taphonomy (continental flooding), the evolution of predators (e.g., sharks, mosasaurs, large theropods) and the Taxophile Effect.
The fossil record of trace fossils provides substantial information about the evolution of vertebrate feeding. In addition, relevant vertebrate and invertebrate ichnofossils provide significant insight into major evolutionary events. The earliest evidence of predation is from terminal Neoproterozoic trace fossils. The earliest definitive vertebrate feeding traces are spiral coprolites and regurgitalites from the Late Ordovician, but earlier bromalites could pertain to vertebrates. The oldest dentalite is from the Middle Silurian. Consumulites occur in Middle Silurian fish, but the first recognizable contents are from the Early Devonian. Bromalites and dentalites provide evidence for major evolutionary events including the Great Ordovician Biodiversification Event, Middle Paleozoic Marine Revolution, and Mesozoic Marine Revolution, as well as various proposed mass extinctions, real and imagined.
3
Consumulites are important in understanding the evolution of the avian digestive Bauplan. We also introduce the following terms: (1) cropalite for preserved contents of the crop, (2) proventrilite for contents of the proventriculus; (3) aspirationalite (from the medical term for food in the airway) for consumulites that preserve prey in the oral cavity with a significant portion extending exteriorly; (4) dislocational evisceralite for a portion of the gastrointestinal tract is physically removed from a carcass; and (5) preservational evisceralite for preferential fossilization of the gastrointestinal tract. There is an almost 200-year history of naming vertebrate tracks, but there is an unjustified reluctance to apply a binominal ichotaxonomy to traces related to feeding that impedes the development of their study.
Nimravids were the first carnivorans to evolve saberteeth, but previously portrayed as having a narrow evolutionary trajectory of increasing degrees of sabertooth specialization. Here I present a novel hypothesis about the evolution of this group, including a description of Eusmilus adelos, the largest known hoplophonine, which forces a re-evaluation of not only their relationships, but perceived paleoecology. Using a tip-dated Bayesian analysis with sophisticated evolutionary models, nimravids can now be viewed as following two paths of evolution: one led to numerous early dirk-tooth forms, including E. adelos, while the other converged on living feline morphology, tens of millions of years before its appearance in felids.
Motivation
Body mass is one of the most important determinants of animal ecology. Unlike other important traits it is also readily inferable from fossils and it is therefore one of the only traits that can be directly analysed and compared between fossil and contemporary communities. Despite this, no comprehensive database of the body mass of larger clades of extinct species exists. Analysis of fossils has therefore been restricted to small clades or to smaller, potentially biased, subsets of species. We here describe CarniFoss, an open-access database of body masses of all 1,322 extinct species of non-pinniped Carnivoramorpha and two related extinct groups of carnivorous mammals, Hyaenodonta and Oxyaenidae.
Main types of variables contained
We gathered lengths of teeth of fossil and extant species and body mass for extant species and a few of the best-known fossil species. Following this we estimated body mass for all species through phylogenetic imputation.
Spatial location and grain
Global, terrestrial.
Time period and grain
We collected data on all known species within the focal groups. The known species all lived in the Palaeogene, Neogene or Quaternary (i.e., the last 66 Myr).
Major taxa and level of measurement
We searched for data on reported tooth size of all described species of Carnivoramorpha (excluding pinnipeds) and selected extinct related groups (Hyaenodonta and Oxyaenidae). We combined this with measured body mass for all extant species and inferred body mass based on long-bones for selected extinct species, as well as a species-level phylogeny including all extant and extinct species in the group, and inferred the body mass for all species using phylogenetic imputation.
Software format
Data are provided as a series of .csv files, with all metadata in a separate PDF file.
A paleoparasitological analysis was carried out on a large coprolite assigned to a carnivoran mammal, recovered from the Municipality of Uruguaiana, in the western region of the State of Rio Grande do Sul, Brazil, where the Upper Pleistocene Touro Passo Formation crops out. For this, an individual sample was extracted from the specimen using an electric drill, dissociated with 10% hydrochloric acid solution, washed with distilled water, and sifted through a 500 mesh Tyler sieve. After laboratory processing, the sediment retained on the sieve was mixed with glycerin and examined by optical microscopy, which revealed the presence of 14 protozoan oocysts and three nematode eggs. The morphological characteristics of the oocysts (i.e., spherical shape, thick-walled, internal zygote apparently at the beginning of sporulation, as well as their size) and of the eggs (i.e., ovoidal shape, rounded ends, smooth surface, thin-shelled, embryo in their interior, along with their morphometry) suggest that these specimens belong respectively to the orders Eucoccidiorida and Strongylida (Family Ancylostomatidae) represented by several parasitic species of the alimentary tract of modern carnivore. This is the first record of paleoparasites discovered in a vertebrate host from the Touro Passo Formation.
Palaeo-wildfire, which had an important impact on the end Permian terrestrial ecosystems, became more intense in the latest Permian globally, evidenced by extensive occurrence of fossil charcoals. In this study, we report abundant charcoals from the upper part of the Xuanwei Formation and the Permian–Triassic transitional Kayitou Formation in the Lengqinggou section, western Guizhou Province, Southwest China. These charcoals are well-preserved with anatomical structures and can be classified into seven distinctive types according to their characteristics. Organic carbon isotopic analyses of both bulk rocks and charcoals show that the δ¹³Corg values in the Kayitou Formation are notably more negative than those in the Xuanwei Formation, with a negative excursion of 4.08‰ immediately above the volcanic ash bed in the middle of the uppermost coal bed of the Xuanwei Formation. Charcoals with high reflectance values (Romean = 2.38%) are discovered below the ash bed. By contrast, the reflectance values (Romean = 1.51%) of the charcoals in the Kayitou Formation are much lower than those of the Xuanwei Formation, indicating the palaeo-wildfire types have changed from crown fires to surface fires, which was probably due to the retrogression of vegetation systems during the extinction. Based on the above evidence, we suppose that palaeo-wildfires became more frequent and more severe since the climate became drier during the latest Permian in Southwest China, and the eventual vegetation changeover of the terrestrial ecosystems in Southwest China could be caused by volcanism.
In the southern cone of South America, inhabit a large diversity of Neotropical carnivores. Carnivore coprolites are a valuable source of paleoecological and paleoparasitological information. The rock shelter Gruta del Indio (GI) is an emblematic archeological and paleontological site located from Mendoza, Argentina. Several studies were conducted at this site, which provided a stratified sequence spanning the last ∼31 ky BP. The aim of this work was to study parasite remains found in coprolites assigned to carnivores from GI, with the purpose of contributing to the paleoecological knowledge of the site. Twenty coprolites were examined for parasites. Samples were rehydrated in a 0.5% water solution of trisodium phosphate, then homogenized, filtered, and processed by spontaneous sedimentation. The macroscopic remains were separated and dried at room temperature and were examined for diet analysis. All micromammal prey belonged to the Order Rodentia, and six histricomorphs could be identified. Also, bird bones, plant, arthropod, and hair remains were found. Five coprolites contained parasite remains, and eight nematode species were recovered. This is the first paleoparasitological study at this site, and the findings broaden the knowledge of the biogeographic history of the gastrointestinal helminths found. The obtained results evidence the importance of carnivore coprolite studies recovered from archeological contexts in the reconstruction of paleoecological scenarios. Also, the importance of carnivores for the dispersion of their own parasites and the parasites of their prey are discussed. These parasitological findings contribute with the study of the presence of potential parasitic zoonoses in the Holocene.
Borophagine canids, such as Borophagus, and bone-cracking hyaenids, share several craniodental features interpreted as adaptations to a durophagous diet. There are possible coprolites of Borophagus from the Miocene of Texas. Well preserved specimens of Borophagus coprolites occur in the late Miocene of California. A new specimen from the Pliocene of New Mexico also represents a coprolite of Borophagus. Borocopros wangi igen. et isp. nov. is the segmented coprolite of Borophagus from the Miocene and Pliocene of western North America. The derived borophagine Borophagus may have been the most efficient bone-cracking member of the sub-family. Localities yielding borophagines should be investigated for coprolites. The ichnotaxon indicates the presence of Borophagus, even in the absence of osteological specimens. There are two biogeographic and taphonomic provinces of vertebrate coprolites during the late Cenozoic: (1) Castrocopros province of the New World characterized by a dominance of herbivore coprolites, but also borophagine coprolites; and (2) Hyaenacoprus province in the Old World, which is dominated by hyena coprolites.
The time scale for marine magnetic anomalies for the Late Cretaceous through Neogene (C-sequence) and Middle Jurassic through Early Cretaceous (M-sequence with deep-tow extension) has been calibrated through magnetostratigraphic studies to biostratigraphy, cycle-stratigraphy and selected radioisotope-dated levels. The majority of the geomagnetic polarity time scale for the past 160 Myr is constructed by fitting spreading rate models to these constraints. The status of the geomagnetic polarity time scale for each geologic period is summarized in the appropriate period chapters.
This chapter incorporates some of the discussions from the Geomagnetic Polarity Time Scale chapters in GTS2004 and GTS2012 by J.G. Ogg and A.G. Smith. The derivation of age-models for the C-sequence and the M-sequence of polarity chrons are new to GTS2020.
Fossil records of vertebrate integuments are relatively common in both rocks, as compressions, and amber, as inclusions. The integument remains, mainly the Mesozoic ones, are of great interest due to the panoply of palaeobiological information they can provide. We describe two Spanish Cretaceous amber pieces that are of taphonomic importance, one bearing avian dinosaur feather remains and the other, mammalian hair. The preserved feather remains originated from an avian dinosaur resting in contact with a stalactite-shaped resin emission for the time it took for the fresh resin to harden. The second piece shows three hair strands recorded on a surface of desiccation, with the characteristic scale pattern exceptionally well preserved and the strands aligned together, which can be considered
the record of a tuft. These assemblages were recorded through a rare biostratinomic process we call “pull off vestiture” that is different from the typical resin entrapment and embedding of organisms and biological remains, and unique to resins. The peculiarity of this process is supported by actualistic observations using sticky traps in Madagascar. Lastly, we reinterpret some exceptional records from the literature in the light of that process, thus bringing new insight to the taphonomic and palaeoecological understanding of the circumstances of their origins.
Coprolites (fossilized faeces) can provide valuable insights into species' diet and related habits. In archaeozoological contexts, they are a potential source of information on human-animal interactions as well as human and animal subsistence. However, despite a broad discussion on coprolites in archaeology, such finds are rarely subject to detailed examination by researchers, perhaps due to the destructive nature of traditional analytical methods. Here, we have examined coprolitic remains from the Neolithic (third millennium BCE) settlement at Skara Brae, Orkney, using a range of modern methods: X-ray computed tomography, scanning electron microscopy, lipid and protein analysis (shotgun proteomics of the coprolite matrix as well as collagen peptide mass fingerprinting of isolated bone fragments). This combined approach minimised destructiveness of sampling, leaving sufficient material for subsequent study, while providing more information than traditional morphological examination alone. Based on gross visual examination, coprolites were predominantly attributed to domestic dogs (Canis familiaris), with morphologically identified bone inclusions derived from domestic sheep (Ovis aries) and common voles (Microtus arvalis). Partial dissection of a coprolite provided bone samples containing protein markers akin to those of domestic sheep. Considering the predominance of vertebral and distal limb bone fragments, Skara Brae dogs were probably consuming human butchery or meal refuse, either routinely fed to them or scavenged. The presumably opportunistic consumption of rodents may also have played a role in pest control.
The exceptional fossil site of Las Hoyas (upper Barremian, Cuenca, Spain) yields abundant small to medium vertebrate coprolites, hindering the search for parasites. We studied the contents of 29 coprolites that were previously classified into distinct morphotypes. Several parasitic eggs were retrieved from two of these coprolites, confirming the second record of digenea trematode eggs and nematode (ascaridid) eggs from an Early Cretaceous locality. The cylindrical coprolite containing anisakid eggs was likely produced by a crocodylomorph as the parasite host, whereas the bump-headed lace coprolite indicates the role of a fish as an intermediary or definitive host of the trematodes and ascaridids. These trace and body fossils show that the Las Hoyas 126-129 Ma lacustrine ecosystem documents the early connection between basal Gonorynchiformes fish and digenetic trematodes.
The fossil record of parasitic helminths is often stated to be severely limited. Many studies have therefore used host constraints to constrain molecular divergence time estimates of helminths. Here we review direct fossil evidence fossil evidence for several of these parasitic lineages belong to various phyla (Acanthocephala, Annelida, Arthropoda, Nematoda, Nematomorpha, Platyhelminthes). Our compilation shows that the fossil record of soft-bodied hel-minths is patchy, but more diverse than commonly assumed. The fossil record provides evidence that ectoparasitic helminths (e.g., pentastomids) have been around since the early Paleozoic, while endoparasitic helminths arose at least during, or possibly even before the late Paleozoic. Nematode lineages parasitizing terrestrial plant and animal hosts have been in existence at least since the Devonian and Triassic, respectively. All major phyla (Acanthocephala, Annelida, Platyhelminthes. Nematoda, Nematomorpha) had evolved endoparasitic lineages at least since the Mesozoic. Interestingly, although parasitism is considered derived within Metazoa, the oldest evidence for Nematoda and Platyhelminthes includes body fossils of parasitic representatives. Furthermore, the oldest fossil evidence of these parasitic lin-eages often falls within molecular divergence time based on host co-evolution suggesting the fossil record of helminths themselves might be just as good or at least complementary (and less circular in justification) to calibration based on host associations. Data also provide evidence for obvious host switches or extinctions,
A new hyaenid specimen, a fragmentary mandible with p3-4 from the Late Miocene (MN 9/10) of Eppelsheim, is described and discussed. It is assigned to Protictitherium crassum (Filhol, 1883) based on its morphology, especially low mandible height, slenderness of premolars, and height of distal accessory cuspids in p4. P. crassum is a well-known, temporally and spatially widespread Middle and Late Miocene hyaenid carnivoran. The analysis of the new specimen allows the re-assignment of another mandible fragment from Eppelsheim to P. crassum which was described as "Ictitherium robustum" by Koenigswald (1928). Though both specimens represent the upper dental size range of the species, their morphology and small size exclude other hyaenid taxa, such as Ictitherium viverrinum or Thalassictis robusta. Very large specimens of P. crassum were previously described as separate species (P. arambourgi), but this taxon is now thought to be a synonym of P. crassum. The same is true for "Ictitherium" tauricum, which is also similar in size and morphology, but might be synonymous with P. crassum, as well. Protictitherium crassum remains the only hyaenid taxon known from the Dinotheriensande.
Coprolites are a useful tool to obtain information related to the life history of the producer, trophic interactions, biodiversity, paleoenvironments, and paleoclimate, among other issues. We analyze here a sample of 111 coprolites recovered from levels of the Santa Cruz Formation (lower–middle Miocene, Burdigalian–early Langhian), outcropping in different localities of the Santa Cruz Province, Patagonian Argentina. Based on size and shape, two morphotypes were identified: coprolites assigned to morphotype I vary from ovoid to subspherical in shape, while coprolites assigned to morphotype II are cylindrical in shape. Several coprolites have bone and teeth inclusions belonging to small mammals (i.e., Octodontoidea and/or Chinchilloidea rodents). Morphometry, composition, and taphonomy of the bone remains suggest that the coprolites were produced by carnivorous mammals. According to the features of the guild of carnivorous mammals from the Santa Cruz Formation, we interpret that hathliacynids and/or small borhyaenoids (Sparassodonta) are the most probable producers. Different traces recorded in the coprolites, such as borings and putative eggs, suggest that the feces were exploited by coprophagous insects, probably dung beetles, for different purposes such as feeding and possible oviposition.
A total of 462 coprolites from three localities exposing Upper Cretaceous deposits in the Münster Basin, northwestern Germany, have been subjected to an array of analytical techniques, with the aim of elucidating ancient trophic structures and predator-prey interactions. The phosphatic composition, frequent bone inclusions, size and morphology collectively suggest that most, if not all, coprolites were produced by carnivo-rous (predatory or scavenging) vertebrates. The bone inclusions further indicate that the coprolite producers preyed principally upon fish. Putative host animals include bony fish, sharks and marine reptiles-all of which have been previously recorded from the Münster Basin. The presence of borings and other traces on several coprolites implies handling by coprophagous organisms. Remains of epibionts are also common, most of which have been identified as the encrusting bivalve Atreta. Palynological analyses of both the coprolites and host rocks reveal a sparse assemblage dominated by typical Late Cretaceous dinoflagellates, and with subordinate fern spores, conifer pollen grains and angiosperm pollen grains. The dinoflagellate key taxon Exochosphaeridium cenomaniense corroborates a Cenomanian age for the Plenus Marl, from which most studied coprolites derive. The findings of this study highlight the potential of a multi-proxy approach when it comes to unravelling the origin, composition and importance of coprolites in palaeoecosystem analyses.
We analyze the multivariate pattern of lower and upper cheek dentition for the family Hyaenidae along its evolutionary history. A total of 11,698 individual measurements of lengths and widths for the main postcanine teeth were collected for 54 extinct and three extant species of this family and analyzed by means of principal component analyses. Our results indicate that the functional aspects are better reflected by lower cheek dentition as a result of mosaic evolution. The multivariate structure captured by the three first principal components correspond to different adaptive strategies. The two first components characterize the main groups of ecomorphs, while hunting species separate from scavengers along the third axis. In the context of Hyaenidae, the post-canine cheek dentition of Parahyaena brunnea and Hyaena hyaena shows an extreme degree of specialization in scavenging.
– A paleoparasitological investigation was carried out on Crocodyliformes coprolites discovered in the Municipality of Santo Anastácio, in the southwestern region of the State of São Paulo. Individual samples were extracted from the surface and internal portion of each coprolite which revealed the presence of thick-shelled parasite eggs in three of them. The morphological characteristics of the specimens, i.e., rounded or oval shape, thick-shelled with mammillated surface, as well as their size, suggest that these eggs belong to the superfamily Ascaridoidea, which are gastrointestinal parasite nematodes of vertebrates, popularly known as “roundworms”. This is the first record of Ascaridoidea eggs discovered in Crocodyliformes hosts from the Upper Cretaceous of Brazil. © 2018, Sociedade Brasileira de Paleontologia. All rights reserved.
This paper presents a study of the macromammalian fauna recovered from Mousterian levels of Llonin Cave. The sample is highly heterogeneous and comprises six species of ungulates, including Rupicapra pyrenaica, Capra pyrenaica, and Cervus elaphus, and seven species of carnivores, predominantly Ursus spelaeus, Crocuta spelaea, Canis/Cuon and Panthera pardus. The archaeozoological and taphonomic study of the remains shows preferential use of basal levels of the cave as a den for hyenas and leopards. Neanderthals were also present during this phase and they would have acted mainly on deer and some caprines, while the action of hyenas would mainly have been linked to scavenging of elements left by humans and the introduction of bear remains. Leopards would have transported caprines in order to consume them. The study of several coprolites confirms that hyenas and leopards were the main occupants of the cave. The information from the animals processed by humans together with other archaeological evidence and the intervention of various carnivores in these basal levels enables us to characterise a palimpsest of occupations that would have been short and sporadic in the case of humans.
Depositional sequences originating in semi-enclosed basins with endemic biota, partly or completely isolated from the open ocean, frequently do not allow biostratigraphic correlations with the standard geological time scale (GTS). The Miocene stages of the Central Paratethys represent regional chronostratigraphic units that were defined in type sections mostly on the basis of biostratigraphic criteria. The lack of accurate dating makes correlation within and between basins of this area and at global scales difficult. Although new geochronological estimates increasingly constrain the age of stage boundaries in the Paratethys, such estimates can be misleading if they do not account for diachronous boundaries between lithostratigraphic formations and for forward smearing of first appearances of index species (Signor-Lipps effect), and if they are extrapolated to whole basins. Here, we argue that (1) geochronological estimates of stage boundaries need to be based on sections with high completeness and high sediment accumulation rates, and (2) that the boundaries should preferentially correspond to conditions with sufficient marine connectivity between the Paratethys and the open ocean. The differences between the timing of origination of a given species in the source area and timing of its immigration to the Paratethys basins should be minimized during such intervals. Here, we draw attention to the definition of the Central Paratethys regional time scale, its modifications, and its present-day validity. We suggest that the regional time scale should be adjusted so that stage boundaries reflect local and regional geodynamic processes as well as the opening and closing of marine gateways. The role of eustatic sea level changes and geodynamic processes in determining the gateway formation needs to be rigorously evaluated with geochronological data and spatially-explicit biostratigraphic data so that their effects can be disentangled.
Borophagine canids have long been hypothesized to be North American ecological ‘avatars’ of living hyenas in Africa and Asia, but direct fossil evidence of hyena-like bone consumption is hitherto unknown. We report rare coprolites (fossilized feces) of Borophagus parvus from the late Miocene of California and, for the first time, describe unambiguous evidence that these predatory canids ingested large amounts of bone. Surface morphology, micro-CT analyses, and contextual information reveal (1) droppings in concentrations signifying scent-marking behavior, similar to latrines used by living social carnivorans; (2) routine consumption of skeletons; (3) undissolved bones inside coprolites indicating gastrointestinal similarity to modern striped and brown hyenas; (4) B. parvus body weight of ~24 kg, reaching sizes of obligatory large-prey hunters; and (5) prey size ranging ~35–100 kg. This combination of traits suggests that bone-crushing Borophagus potentially hunted in collaborative social groups and occupied a niche no longer present in North American ecosystems.
This open access book offers a fully illustrated compendium of glossary terms and basic principles in the field of palynology, making it an indispensable tool for all palynologists. It is a revised and extended edition of “Pollen Terminology. An illustrated handbook,” published in 2009. This second edition, titled “Illustrated Pollen Terminology” shares additional insights into new and stunning aspects of palynology. In this context, the general chapters have been critically revised, expanded and restructured. The chapter “Misinterpretations in Palynology” has been extended with new research data and additional ambiguous terms, e.g., polyads vs. massulae; the chapter “Methods in Palynology” has been extensively enhanced with illustrated protocols showing the majority of the methods and techniques used when studying recent and fossil pollen with LM, SEM and TEM. Moreover, additional information about the description and publication of pollen data is provided in the chapter “How to Describe and Illustrate Pollen Grains.” Various other parts of the general chapters have now been updated and/or extended with more comprehensive textual passages and new illustrations. The chapter “Illustrated Pollen Terms” now features new and more appropriate examples of each term, including additional LM micrographs. Where necessary, the entries for selected pollen terms have been refined by rewording or adding definitions, illustrations, and new micrographs. Lastly, new terms are included, such as “suprasculpture” and the prefix “nano-“ for ornamentation features.
The chapter “Illustrated Pollen Terms” is the main part of this book and comprises more than 300 widely used terms illustrated with over 1,000 high-quality images. It provides a detailed survey of the manifold ornamentation and structures of pollen, and offers essential insights into their stunning beauty.
Springer link:
http://www.springer.com/de/book/9783319713649
The macro-and microscopic structure of the mammal hair is amazingly diverse, being specific and appropriate to identify the species, genera or higher taxa. The identification of mammals using their hair morphological characters requires time and practice, so this method is underproportioned in the era of molecular investigations. At the same time, the hair is the most frequent and often the only life sign of mammals in the field, moreover, the most recent biodiversity and conservation biology studies prefer the use of non-invasive methods, including hair-trapping, bird-nest analysis, and the identification of the hairs as remnants of prey taxa. The trichomorphology can support the taxonomic work, the understanding of the phylogenetic relationships, the adaptation to the environment, and the forensic work. The reliability of the identification of mammals based on their hair characters depends strongly on the reliability of information content of hair characters, so the aim of the author is to help these studies summarizing her many years of experience in this field.
The book encloses a resume on the history of trichomorphology, the anatomy of mammal hair, the most important physiological and taxonomical aspects. The nomenclature and a system of the patterns of hair is a crucial trial to establish an etalon for the further studies. The Atlas chapter contains the descriptions of 123 mammal species occurring in Central Europe. Several hundreds of illustrations and the identification key to the guard hairs are provided for the users to promote their activity.
The book is addressed mainly to the potential users, the scholars and the experts dealing with trichomorphology: biologists, wildlife biologists, conservation biologists, hunters, palaeontologists, archaeologists, wildlife forensic experts. Hopefully, it will provide the enjoyment of discovering this micro-world for the inquiring naturalists and readers, too.
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).
Microstructural post mortem changes to skeletal tissues by microorganisms are driven by several factors including the death history of an animal, its decomposition trajectory, and the depositing environment itself. The study we describe here brings together material from recent and fossil contexts that are depositionally distinct from a terrestrial-marine transitionary shoreline environment. We compare these changes with those of marine environments previously identified in the Mary Rose material, and those of continental waters (lakes) previously identified in the Cerro de la Garita (Concud) site, and we document this against bacterially related changes observed from terrestrial contexts. A new microstructural change identified in material from terrestrial sites is also described relating to rootlet damage. By considering microstructural change in skeletal tissues, it is maybe possible to ascribe environmental context, or, to better understand the complexity of material presented by transitionary environments.
The fossil record of exceptionally preserved soft tissues in Konservat- Lagerstätten provides rare yet significant insight into past behaviours and ecologies. Such deposits are known to occur in bursts rather than evenly through time, but reasons for this pattern and implications for the origins of novel structures remain unclear. Previous assessments of these records focused on marine environments preserving chemically heterogeneous tissues from across animals. Here, we investigate the preservation of skin and keratinous integumentary structures in land-dwelling vertebrates (tetrapods) through time, and in distinct terrestrial and marine depositional environments. We also evaluate previously proposed biotic and abiotic controls on the distribution of 143 tetrapod Konservat-Lagerstätten from the Permian to the Pleistocene in a multivariate framework. Gap analyses taking into account sampling intensity and distribution indicate that feathers probably evolved close to their first appearance in the fossil record. By contrast, hair and archosaur filaments areweakly sampled (five times less common than feathers), and their origins may significantly pre-date earliest known occurrences in the fossil record. This work suggests that among-integument variation in preservation can bias the reconstructed first origins of integumentary novelties and has implications for predicting where, and in what depositional environments, to expect further discoveries of exquisitely preserved tetrapod integument. © 2017 The Author(s) Published by the Royal Society. All rights reserved.
The present Thesis is focused on the study of the carnivorans that have been discovered in the hominid locality of Hammerschmiede. The age of the locality is at the base of Late Miocene and it is situated in Bavaria, Germany. Five articles have already been conducted based on this material, while more specimens are presented here accompanied by a discussion concerning the carnivoran guild of the locality.
The main part of this Thesis includes three introductory chapters. The first chapter deals with the order Carnivora. The phylogenetic relationships of this group are discussed, followed by an extensive presentation of the representatives of this order during the Miocene of Europe. This part was considered essential, because of the extreme diversity of the Hammerschmiede carnivorans that made required a deep understanding of the current knowledge on this group. The second part of the introduction deals with some characteristics of the Miocene of Europe. The carnivoran guilds of several localities are studied, followed by some remarks on palaeogeography and palaeoclimatology. Finally, the last part of the introduction has to do with the locality of Hammerschmiede. The most updated data for the faunal, floral and abiotic components of the locality are demonstrated, together with a historical summary of the studies concerning Hammerschmiede.
The chapter 4 of material and methods includes a detailed table containing the material used in this project, as well as the methodologies used for its study (dental nomenclature, anatomical nomenclature, measurements, guild analysis).
The next part (chapter 5) of the Thesis includes the five articles that have already been conducted for peer-review academic journals. The first paper consists of a taxonomic, biostratigraphic and palaeoecologic review of the genus Semigenetta (Viverridae), reporting some material of Semigenetta sansaniensis and Semigenetta grandis from Hammerschmiede. The second paper reports the presence of a new species of otter, Vishnuonyx neptuni (Mustelidae), accompanied by palaeogeographical and palaeoecological notes. The third paper concerns the material of hyenas (Hyaenidae) found in the locality, presenting dental material of the ictithere Thalassictis montadai, together with a large bone-cracking hyaenid, as well as a biostratigraphic review of relevant forms in Europe. The fourth paper presents the extreme diversity of the small carnivoran forms found in Hammerschmiede, together with a preliminary palaeoecological comparison. The discovered species include: “Martes” sansaniensis, “Martes” cf. munki, “Martes” sp., Circamustela hartmanni (a new species for this genus), Laphyctis mustelinus, Guloninae indet., Eomellivora moralesi, Vishnuonyx neptuni, Lartetictis cf. dubia, Paralutra jaegeri, Trocharion albanense, Palaeomeles pachecoi, Proputorius sansaniensis, Proputorius pusillus, Alopecocyon goeriachensis, Simocyoninae indet., Potamotherium sp., Semigenetta sansaniensis, Semigenetta grandis and Viverrictis modica. Finally, the fifth paper concerns material of the primitive giant panda Kretzoiarctos beatrix (Ursidae) accompanied with a diet estimation for this species based on dental texture microwear analysis.
More material, which either was discovered after the publications or was considered to consist of a separate subject, was studied for the families: Amphicyonidae (1 species), Ailuridae (more material of Alopecocyon goeriachensis), Mephitidae (more material of Proputorius sansaniensis), Mustelidae (more material of Paralutra jaegeri and Lartetictis cf. dubia), Phocidae (1 species), Felidae (Pseudaelurus quadridentatus and Metailurini indet.), Barbourofelidae (1 species) and Hyaenidae (more material of Thalassictis montadai). Additionally, a preliminary description of the coprolites found in the locality has been attempted.
Consequently, the carnivoran guild of the locality includes 28 species. This makes Hammerschmiede the third most diverse locality in the Miocene of Europe (surpassed only by the fissure-fillings Wintershof-West and La Grive-Saint-Alban). This is especially impressive, based on the relatively low number of identifiable specimens (n=122), as also shown in rarefaction analysis. The HAM 5 layer alone has yielded 21 species, in comparison to the 15 species found in HAM 4. A biostratigraphic analysis demonstrated that the locality includes mostly Aragonian, but also some Vallesian forms, including several First Occurrence Dates and Last Occurrence Dates. Species diversity for the discovered families reveals that the profile seen in Hammerschmiede does not resemble that of any other Miocene locality of Europe. A detailed attribution of all the discovered species to the available categories for Dietary Habits, Locomotor Lifestyle and Body Mass has been presented. Quantitative analysis of these data show that the locality mostly includes small- to medium-sized carnivorans, especially in HAM 4. Additionally, many species are scansorial or semi-aquatic, proving the strong influence of the forested river in the locality. Palaeoecological comparison of the discovered species (through cluster analysis and 2D+ plots) shows that most species are able to coexist without competition, whereas other were found to occupy very similar niches.
Concluding, the locality of Hammerschmiede has proven to conceal an astonishing diversity of mammalian carnivores. Based on the presented data, it is sure that it will be a reference locality for the study of the Miocene carnivorans of Europe in terms of taxonomy, biostratigraphy and palaeoecology. Some possible future objectives are mentioned at the end of the discussion.
The present paper deals with new hyaenid material from the locality of Hammerschmiede (Bavaria, Germany). The described specimens are attributed to two forms: most of the specimens belong to the species Thalassictis montadai, whereas one I3 is attributed to a large bone-cracking hyena. The material comes from the layers HAM 5 (11.62 Ma) and HAM 6 (slightly younger than 11.44 Ma) of Hammerschmiede (base of Late Miocene). The species Thalassictis montadai is well-known from late Aragonian and early Vallesian localities of central and southern Europe and west Asia. The presented material enables us to make a short review of the state-of-the-art about the fossil record of this species and to discuss its intraspecific variability. A gradual replacement of Thalassictis montadai, Thalassictis robusta and Hyaenictitherium wongii in Europe is demonstrated, until the arrival of canids during the latest Miocene. Additionally, the upper incisor of the large hyaenid creates some interesting questions concerning the first appearance of the crocutoid hyenas in the fossil record and their dominance over the percrocutoids.
Coprophagy is defined as the consumption of one’s own faeces (autocoprophagy) or the faeces of other individuals of the same or other species (allocoprophagy). The consumption of one’s own faeces or the faeces of conspecifics is common, in particular for rodents and lagomorphs. However, the consumption of faeces of individuals of another species has rarely been described for vertebrates. In this study, we describe occurrence of coprophagy of African wild dog faeces by hooded vultures and spotted hyaenas in Mana Pools National Park, Zimbabwe. Between September 2017 and November 2020, we radio-tracked six collared African wild dog packs and recorded interactions with spotted hyaenas and hooded vultures. When spotted hyaenas and/or hooded vultures were present, they engaged in coprophagy of African wild dog faeces in 38.5% and 65.5% of the cases respectively. For both species coprophagy was not related to season, drought, time of day or size of the African wild dog pack. Hooded vultures especially, often engaged in coprophagy when they accompanied African wild dogs while they were resting, suggesting such an association may be intended to have access to faeces. Allocoprophagy in wild vertebrates usually serves as an additional source of energy and/or nutrients. Further research is required to determine the content of African wild dog faeces and the potential nutritional benefits for spotted hyaenas and hooded vultures. However, it is clear from our and other studies that the critically endangered hooded vulture forms close associations with the endangered African wild dog. Such associations may play a role in the hooded vulture's survival and should therefore be considered in the conservation strategy of this species.
We describe two carnivoran coprolites found in the pseudokarst natural carnivore trap of Batallones-3, from the Late Miocene of Spain. The larger one, comprising multiple indistinguishable fragments of broken and corroded bones, indicates that the producer of the dropping might have been highly capable of crushing the softer parts of large bones. On the other hand, the smaller one shows several relatively larger and more complete bone fragments, thus exhibiting a greater capacity to break and swallow large portions of bone. The external morphology of the large coprolite is similar to that of extant bears, whereas the smaller one more closely resembles that of the living insectiv-orous hyaenid Proteles in morphology, on the one hand, and that of the viverrid Gen-etta in size, on the other hand. We hypothesize that the amphicyonid Magerycion anceps was the producer of the large coprolite and the jackal-sized basal hyaenid Protic-titherium crassum excreted the smaller one. Thus, we present the first direct evidence of a bone durophagous diet in the carnivorans of Batallones.
We analyse the evolution of carnivoran guilds (body mass, locomotor pattern, and diet preference) across the Middle to Upper Miocene boundary based on a comparison of the carnivoran fauna from Steinheim (Mammal Neogene zone MN 7/8; Servallian to early Tortonian), and from the Eppelsheim Formation (Mammal Neogene zone MN 9/10; mid-Tortonian), Germany. Results reveal a massive faunal turnover between these two communties due to taxonomic differences up to the family level. Guild structures of both carnivoran faunas were similar, implying that Late Miocene taxa replaced Middle Miocene taxa in their respective niches rather than new ecological strategies being added to the guild. This is not surprising, as (1) the majority of Late Miocene taxa were derived from European Middle Miocene ancestors and mostly share their paleoecology, (2) of the few undisputed Late Miocene immigrant taxa (Eomellivora, Simocyon, Sivaonyx), only Eomellivora adds new ecological strategies to the guild, and (3) both faunas occupied the same kind of partly open woodland paleoenvironment. Our paper supports previous studies, suggesting that ecological diversity remains stable across the Middle to Upper Miocene boundary and that similar environments produce similar carnivoran guild structures irrespective of taxonomical composition
The Eocene Nanjemoy Formation crops out on the Maryland and Virginia Coastal
Plain, along the eastern coast of the United States. This formation is composed of
sands, silts and clays and is divided into the Potapaco and Woodstock members.
Remains of fishes, reptiles, birds, mammals, molluscs, fruits and seeds are common in
the Potapaco Member, in addition to vertebrate coprolites. Here, we present an analysis
of more than 2000 coprolites from the Fisher/Sullivan Site in Virginia. The chemical
composition (phosphatic) and the type of inclusions (fish bones) indicate that only
scats of carnivorous animals were preserved. The analysed specimens were grouped
into six morphotypes: (1) the cylindrical morphotype is a cylinder with rounded ends;
(2) the segmented morphotype is a cylinder segmented with rounded ends, and occasionally
one end is concave; (3) the oval morphotype represents a bean‐shaped coprolite;
(4) the scroll morphotype is cylindrical to conical in lateral view and has coils seen
only at the ends; (5) the folded morphotype is a spiral that is concentrically folded; and
(6) the sinuous morphotype is serpentine, with rounded ends. Coprophagy‐related
scrape traces occur in different morphotypes and represent both invertebrate burrows
and bite traces made by fishes. The mineralogical and chemical analyses indicate an
early precipitation of phosphate and pyrite minerals, probably induced by the microbial
community. All coprolites at the Fisher/Sullivan Site were produced by fishes: carcharhiniform sharks for the scroll morphotype and lamniform sharks, probably the
genus Carcharias, for the folded morphotype; the oval, cylindrical and segmented morphotypes were likely produced by actinopterygian fishes.
Regurgitalites (fossilized regurgitates) can provide insight into the behavioral ecology and physiology of extinct species, but they are rarely reported because they are difficult to identify and distinguish from coprolites. A compact mass of skeletal material from the Owl Rock Member of the Upper Triassic Chinle Formation of Arizona reveals features that identify it as a regurgitalite. Characteristics of the teeth and osteoderms in the specimen indicate that these remains belong to the pseudosuchian archosaur Revueltosaurus. Chemical and microstructural analysis revealed a dearth of gastric etching, the preservation of muscle fibers, and the absence of a phosphatic matrix, indicating that this bone mass is a regurgitalite and not a coprolite. It was probably produced by a phytosaur, rauisuchid, or temnospondyl, all of which occur in the Owl Rock Member. We offer an identification key to assist in distinguishing between different types of digestive remains produced by vertebrate carnivores.
The aims of the present study were: (1) to examine the parasite fauna found in carnivore coprolites from Cueva Huenul 1 archaeological site, located in northern Patagonia (Neuquén Province, Argentina), (2) to evaluate the role of this carnivores in the cycle of zoonotic parasites in the past, (3) to discuss the possible effect of the infections in humans that inhabit the cave during the Quaternary, and (4) to evaluate possible differences in egg measurements among layers. Several coprolites were obtained from layers dated since the Pleistocene-Holocene Transition. Faeces were whole processed, rehydrated, homogenised, and examined via light microscopy. Eggs of parasites were measured and photographed. Coprolites were assigned to a small feline. A total of 11 parasite species were found (8 nematode, 2 trematode, and 1 coccidian species). The paleoparasitological findings of this study display a great richness of parasitic species present in felines from CH1 through the Pleistocene-Holocene transition, some of them zoonotic. Early human societies that discontinuously occupied the site during the time scale studied could have been exposed to these infections. ARTICLE HISTORY
In 1822, William Buckland first recognized Pleistocene vertebrate coprolites, and they are now known from more multiple localities with a global distribution. Carnivore coprolites dominate, and there are two distinct biogeographic and taphonomic provinces for vertebrate coprolites in the Pleistocene. The Castrocopros province of North and South America is characterized by a dominance of herbivore coprolites, which are preserved almost exclusively in caves. The Hyaenacoprus province in Africa, Europe and Asia is dominated by hyena coprolites that occur mainly in caves, but also in other depositional environments. Pacific Islands may represent a third province characterized by dominant bird coprolites from caves, but the known localities are all Holocene. Mammuthocopros allenorum ichnogen. and ichnosp. nov. is a coprolite of Mammuthus columbi from Utah, USA.
Microbial bioerosion, in all its manifestations, is one of the major factors determining the long-term survival of archaeologically and environmentally important artefacts and ecofacts made from mineralised collagen – bones, antler, teeth and ivory. The bone diagenesis literature contains extensive descriptions of different morphologies and classifications of microbial bioerosion – microscopical focal destruction, Wedl tunnels, linear longitudinal, budded and lamellate tunnelling, etc. but the causative agents remain to be discovered. Palaeontologists are in a similar situation when describing ichnofossils where bioerosion is classified by its characteristic morphology and only tentatively assigned to specific causes. In archaeological bones, Wedl tunnels have traditionally been ascribed to fungi but a re-examination of Wedl's original paper and subsequent literature has shown that all the early specimens examined came from aquatic environments and that euendolithic microflora (cyanobacteria or chlorophytes) might have been responsible. These microorganisms are known to tunnel into marine shells. Linear longitudinal, budded and lamellate tunnelling have all been ascribed to bacteria. However, analyses using a combination of backscatter SEM (BSEM) and mercury intrusion porosimetry (HgIP) of bones excavated from terrestrial soils suggest that the various tunnelling morphologies described by earlier researchers are actually all manifestations of a single architecture, the differences arising from the inherent variability of bone microstructure and the hydrology of the burial environment. An examination of the BSEM and HgIP data also indicates that the bacteria responsible may spread through dead bone tissues by expanding the canalicular network (or dentinal tubules in teeth) rather than creating new tunnels.
Long-term field burial experiments using de-fleshed cow bone specimens have now demonstrated that bacterial tunnelling develops over decades rather than months as was previously thought, even in warm tropical soils. This has obvious implications for the origin of the bacteria responsible (soil bacteria versus endogenous gut bacteria) since the bone specimens were quickly isolated from gut contents. Although no attempt is made here to identify a specific organism or organisms responsible for tunnelling in terrestrial archaeological bones a tentative model is proposed for how cycles of wetting and drying, or changes in local dissolved oxygen levels, could lead a common gram positive soil bacterium to produce the pattern of bacterial bioerosion seen in exhumed bones. This soil organism may act in a similar way to Staphylococcus aureus which proliferates in living bone (causing osteomyelitis) by infiltrating and enlarging the canalicular network.
Lophocyonids are one of the more enigmatic groups of Carnivora in the European Miocene fossil record. Lophocyonids are clearly distinguished from other Feliformia by their peculiar lophodont dental morphology. For this reason, the systematic relationships of the family have been controversial. Here we describe and interpret dental remains from the early Miocene of Sabuncubeli, Turkey, which we attribute to a new genus and species: Izmirictis cani. The phylogenetic analysis allows us to include Izmirictis within a monophyletic group, Lophocyonidae new rank, characterized by the molarization of the anterior premolars (P3 and p4), the lophodont adaptation of the molar dentition and the complex morphology of the incisors. The phylogenetic analysis shows a close relationship between Izmirictis and primitive hyaenids (represented herein by Protictitherium). The divergence between Lophocyonidae and Hyaenidae is estimated by biochronological data to have occurred during the early Miocene (MN2). Dental microwear analysis, although limited by poor dental enamel preservation, indicates that the pronounced lophodonty in Izmirictis cani could be connected to a herbivorous feeding habit.
http://zoobank.org/urn:lsid:zoobank.org:pub:8D75382D-6C22-41C1-A47F-BA3EAF766881
The St. Stefan clay-pit at Gratkorn, at the north-western margin of the Styrian Basin, is an extraordinary fossil locality, due to its diverse and unique plant and animal records. The age of the plant-bearing sediment is Sarmatian, late Middle Miocene. Macroflora - both leaves and fruits - is rich in the locality. This paper provides a taxonomic work of the leaf flora as well as gives a summary of the flora, vegetation and climate. Aquatic vegetation is composed of Ceratophyllum schrotzburgense Hantke, Caldesia europaea Hably sp. nov., which is the first leaf record of Caldesia in Europe, Stratiotes sp., Hydrochariphyllum kvacekii Hably sp. nov., Hydrocharitaceae gen. et sp. indet, Potamogeton martinianus Sitar and Monocotyledonae gen. et sp. indet. In wetland habitats the following taxa existed: Pteris oeningensis Unger, Podocarpium podocarpum (A.Braun) Herendeen, Myrica lignitum (Unger) Saporta, Ulmus carpinoides Göppert emend. Menzel, cf. Alnus sp., Salix varians Göppert, Salix sp. 1., Populus balsamoides Göppert, Decodon sp., Acer integrilobum Weber sensu Walther, Asclepiadaceae vel Cucurbitaceae gen. et sp. indet. The zonal vegetation is represented by Buxus pliocenica Saporta et Marion. © 2017 E. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart, Germany.
Pipestone Springs Main Pocket (PSMP) (Renova Formation, Jefferson County, Montana) has yielded an unusually rich concentration of mostly small-bodied Chadronian (late Eocene) mammals. Coprolites are common at PSMP and indicate which taxa and skeletal elements were consumed, as well as provide insight as to which carnivore species were likely to have deposited their feces at the site. Two distinct coprolite groupings were recognized based on differences in diameter, morphology, and abundance of bone. The sample of larger coprolites ranges in diameter from 16-29 mm and lacks a distinct mode. Some of the larger coprolites contain bone, but if present, it is sparse, poorly preserved, and unidentifiable. The sample of smaller coprolites varies in diameter from 4-15 mm and has a prominent mode at 10-11 mm. Many of the smaller coprolites contain abundant bone that is often highly fragmented. Osteoderms, bone ends, or dental elements identified to taxon in smaller coprolites indicate that marsupials, lizards, lagomorphs, and rodents were prey of smaller PSMP carnivores. Most elements identified as lagomorph or rodent probably represent Palaeolagus temnodon and Ischyromys veterior based on their high abundance in surface collections. Undigested bone from disaggregated carnivore feces did contribute to the large number of small dentigerous elements recovered from surface collections at PSMP, but it probably was not a major source because only two smaller coprolites had an exposed partial dentary or maxilla. The remains of mammalian carnivores are comparatively rare at PSMP and nine taxa are described, 52% of which represent Hesperocyon gregarius and 21% Brachyrhynchocyon dodgei. If the percentage of surface collected specimens for each carnivore taxon at PSMP is a reflection of activity at the depositional site, most of the smaller coprolites probably represent Hes. gregarius. To test this hypothesis, the ratios between feces diameter and body mass, as well as average lower first molar, (m1), length and body mass in an extant canid were applied to m1s of Hes. gregarius and the smaller coprolites from PSMP. Results indicate a mass of 3.3 kg for the smaller PSMP carnivore and 2.9 kg for Hes. gregarius from PSMP, both similar to the mass of Hesperocyon based on skeletal material. Also, the mode of smaller diameter coprolites is more prominent than the mode of the diameter of feces from an extant canid, which suggests that most of the smaller PSMP coprolites represent the activity of a single species, in this case Hes. gregarius. Many larger coprolites at PSMP probably represent Br. dodgei based on its high relative abundance, but the lack of a prominent mode for the larger coprolites suggests that one or more other species also contributed to the sample.
Coprolites (fossil feces) provide important paleoecological information, such as diet of the producer, parasite infestation and gut microbiota, which cannot be accessed through body remains. The Upper Miocene Urumaco Formation, in western Venezuela, has a diverse vertebrate fauna composed of ground sloths, rodents, reptiles, fishes and amphibians, as well as coprolites. Here we describe 106 coprolites produced by a range of carnivorous and herbivorous vertebrates, classified into five different morphotypes based on their size and shape, suggesting different producers. Herbivorous morphotypes (M1, M2 and M5) are the most abundant (59%). The most common inclusions in these coprolites are Poaceae remains, although Eudicotyledoneae fragments, bacteria and cyanobacteria filaments were also found. The M1 morphotype has a bullet shape with longitudinal grooves; the M2 morphotype is rounded to oval, with at least one distinctive tapered end; and the M5 morphotype is a short cylinder with a rough surface, and rounded ends. In contrast, the morphotypes M3 and M4 are attributed by us to carnivores and have different inclusions. While M3 is a large cylinder with rounded ends and contains bacteria, nematode eggs and phytoliths, M4 is sausage-shaped and has muscle cells preserved inside. Based on the co-occurring fauna and the morphology of the coprolites, the probable producers were inferred as the following: M1, a rodent; M2, a notoungulate; M3, a crocodilian; M4, an indeterminate carnivore; and M5, a sirenian. The preservation of herbivorous and carnivorous coprolites with bacterial, parasitological and muscle remains in the same formation and even the same outcrop is rare in the geological record. This unique association and preservation suggests early lithification promoted by rapid burial, although the presence of the boring trace Gastrochaenolites in a single specimen indicates some degree of time-averaging.
With high organic content and abundant microbiota, feces are typically only preserved for more than a couple of days, weeks or months after deposition and most never enter the fossil record. The range of depositional conditions favorable to the feces preservation is narrow, but depends on a large number of biotic and abiotic factors. Moreover, because the chemistry and microbiology of feces often promotes mineralization, discriminating between coprolites and some inorganic sedimentary structures is often difficult. We propose a protocol to identify coprolites in the fossil record that encompasses all the criteria previously defined. We then apply this protocol to identify putative coprolites from the lower Carnian (Upper Triassic) Chañares Formation of northwestern Argentina. Using a variety of analytical methods and several different criteria, we were able to identify them as carnivore coprolites. Based on extant analogs, the most probable producer would be a small carnivorous cynodont or archosauriforms, and the prey is likely a tiny therapsid or archosauriform. This is the first detail report of carnivore coprolites from this unit, and provides direct evidence of trophic links in the Chañares ecosystem.