Coprolites

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.

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.

Hitherto unpublished 14 C and 230 The 234 U determinations from Carihuela Cave (Granada province, Andalusia, Spain) raise a possibility of late survival here of Neanderthals and their Mousterian tech-nocomplex into an advanced stage of the Late Pleistocene (MIS-3), when anatomically-modern humans with Upper Palaeolithic toolkits were penetrating the region, and when also several carnivore taxa competed for access to the cave. Previous palaeopalynological studies are reinforced by new pollen analyses of samples extracted from coprolites. The palaeoecological and sedimentological records bear comparison with new data from the Padul peat deposits in the Sierra Nevada, and are in line with the view that there was late persistence of the Mousterian in Granada. There is a pressing need for renewed international multidisciplinary research at Carihuela Cave, with up-to-date lithostratigraphical and dating techniques that can expand on results obtained from fieldwork undertaken by a previous generation of researchers. Carihuela Cave continues to hold out great promise for analysing Neanderthal palaeoecology during the Late Pleistocene up to the appearance in southeastern Iberian Peninsula of anatomically-modern Upper Palaeolithic people, particularly with regard to the earlier phases of the Middle Palaeolithic at the cave which await intensive excavation but apparently extend back in time to the last interglacial period.

In the period between ~42,500 and ~50,000 calendar years ago, represented in the Gruta da Oliveira archeostratigraphy by levels 9-14, Middle Paleolithic humans took shelter in an area of the site located behind a >20 ton boulder fallen at the entrance. Hyena coprolites were also recovered in this area, especially in layers 10-13, where specific spots were recurrently used as latrines, and small fragments of degraded coprolites and digested bone are ubiquitous in micromorphological thin sections. The pollen in the coprolites suggests that hyenas used the site at times when a temperate forest developed outside, while the charcoal, dominated by Scots pine, gives a cold climate signal. Combined, these data suggest a pattern of alternating use of the cave, with hyena scavenging of human-accumulated dry bone but, given the near-absence of skeletal remains and the lack of juveniles, without any denning.

Cova del Gegant (Barcelona, NE Iberian Peninsula) is a Middle Palaeolithic site presenting evidence of carnivore occupations with some sporadic Neanderthal activity. The rapid, fine sedimentation in layer IIIa permitted the preservation of the remains of carnivore activity and an ephemeral combustion area, although lithics are absent. We used a detailed excavation procedure in combination with Fourier transformed infrared spectroscopy, soil micromorphological, zooarchaeological and taphonomical analyses, and coprogenic analysis to evaluate the interactions of humans and carnivores in the same area of the cave.

Shotgun metagenomics applied to archaeological feces (paleofeces) can bring new insights into the composition and functions of human and animal gut microbiota from the past. However, paleofeces often undergo physical distortions in archaeological sediments, making their source species difficult to identify on the basis of fecal morphology or microscopic features alone. Here we present a reproducible and scalable pipeline using both host and microbial DNA to infer the host source of fecal material. We apply this pipeline to newly sequenced archaeological How to cite this article

Animal dung is evaluated here as a tool to reconstruct recent societal and environmental changes. Studies
completed on the macro- and micro-contents from dung deposited in a mountain cave in Catalonia during the
1970–1980s, preceding the socio-economic changes in the area, was supplemented by the testimony of the
last shepherd alive in the area. This information was also compared with evidence from aerial photographs of
the area surrounding the cave taken in 1956 and 2009. Although taphonomic distortion of the dung shape
precluded its assignment to its producers based on morphology, information from the organic remains
reflects the agro-pastoral activities in the mountainous area of the Catalan coast. A lack of calcium
phosphate and abundant phytolith and epidermal remains from wild species indicates that the dung was
excreted by ovicaprines, who sheltered in the cave during spells of adverse weather. These indicators
also signify that the animals fed only on natural vegetation. Insect remains in the dung include fragments
of chafers, dung and ground beetles, bees and ants, and whilst the latter may have foraged in the cave,
the others may have been accidentally ingested by herbivores. The nannofossils and diatoms found in the
dung derive from tertiary crops 4 km away from the cave but within the range of the ovicaprine herd. The
lack of cereal and vegetable remains in the dung is consistent with the ethnographic record that
shepherds and farmers could not afford to supplement the herds’ diet with fodder crops or grain, a
consequence of the low productivity of the local subsistence agriculture. This study further discusses the
socio-economic impacts of the introduction of intensive farming, industrialisation and tourism in Catalonia
during the latter part of the last century and how they are represented in the dung.
Keywords: Animal dung, Phytoliths, Diatoms, Insects

Coprolites (fossil feces) are important sources of evidence of ancient food webs and ecosystems. Actinomycetes are a fundamental component in the decay of organic matter, and serve as catalysts for nutrient cycles. Recently, gas vesicles filled with numerous verrucose colonies of substrate mycelium of an actinomycete were discovered inside a fossilized spiral amphipolar fish coprolite recovered from mid– Permian deposits of Brazil. These colonies are composed of masses of substrate hyphae, some of which are undergoing segmentation. Arising from the colonies are chains of spores separated by narrow, elongate connectives. The fossil actinomycete is described below as Palaeostromatus diairetus gen. et sp. nov. and represents the oldest known actinomycete associated with vertebrate deposits. Since the colonies occur only inside the coprolite, either Palaeostromatus diairetus gen. et sp. nov. was part of the gut flora or it was acquired from a food source. The only other remains in the coprolite are eighteen paleoniscoid fish scales, which suggests that the producer was a carnivorous/omnivorous fish. This is the oldest record of a direct interaction between vertebrates and actinomycetes.

Until now, the oldest known human hair was from a 9000-year-old South American mummy. Here we report fossil hairs of probable human origin that exceed that age by about 200,000 years. The hairs have been discovered in a brown hyaena (Parahyaena brunnea) coprolite from Gladysvale cave in South Africa. The coprolite is part of a hyaena latrine preserved in calcified cave sediment dated between 195,000 and 257,000 years ago. This find supports the hypothesis that hyaenas accumulated some of the early hominin remains found in cave sites, and provides a new source of information on Pleistocene mammals in the Sterkfontein Valley.

Coprolites are commonly identified in the Pleistocene archaeo-palaeontological record. They have often been described as indirect evidence for the presence of carnivores (usually hyenids) during the formation of a depositional sequence. However, coprolites are a much larger source of information that can provide data relating to factors affecting an archaeological assemblage, including its taphonomic history and palaeoecology. In this paper, two fossil coprolite accumulations dating to the late Early Pleistocene are described: Level TD6.1 of Gran Dolina (Sierra de Atapuerca, Burgos) and Unit II of La Mina (Barranc de la Boella, Tarragona). The results highlight the morphological homogeneity of these hyena coprolites, despite a considerable variation in size. The presence of microspherulites is clearly identified in all of the analysed hyena coprolites. Their composition is rich in elements characteristic of bone-origin, such as calcium (Ca) and phosphorus (P). A major difference, however, differentiating the remains from these sites was a higher frequency of bone fragments in the La Mina coprolites, regarding to TD6.1. Pollen, spores, and palynomorphs are scarce in the samples from both sites, which in fact share similar ecological characteristics such as the presence of pine, wild grasses, and coprophilous fungi. Hyena coprolites are relatively easy to differentiate from those of other taxa; however, they show intra-specific similarities in their micro- and macro-morphology and composition, despite their wide size range. The species of hyenids responsible for the coprolites is not possible to identify based exclusively on the size range, and the importance of a multidisciplinary study of the latrines and the coprolites is discussed as a source of palaeoecological and taphonomic information. In this study, coprolites are analysed, compared, and attributed to a hyenid, identified as Crocuta crocuta in the case of TD6.1, but without specific attribution in the case of La Mina.