A tick from a prehistoric Arizona coprolite

Ticks have never been reported in archaeological analyses. Here, we present the discovery of a tick from a coprolite excavated from Antelope Cave in extreme northwest Arizona. Dietary analysis indicates that the coprolite has a human origin. This archaeological occupation is associated with the Ancestral Pueblo culture (Anasazi). This discovery supports previous hypotheses that ticks were a potential source of disease and that ectoparasites were eaten by ancient people.     

Pollen concentration analysis of Ancestral Pueblo dietary variation

Previous coprolite research on the Colorado Plateau has shown that macrofossils are a useful way of statistically demonstrating prehistoric dietary variation of Ancestral Pueblos (Anasazi). Up until now, pollen concentration from human coprolites has not been used for comparative, statistical study. We present here the statistical analysis of pollen concentration values of coprolites from two Ancestral Pueblo sites, Salmon Ruin and Antelope House. The data show that although most pollen types do not show statistically significant variation, there are some types that show how different Ancestral Pueblo populations adapted to plant resources in different environments. The analysis indicates that future work should focus more on pollen concentration analysis of coprolites.     

Cultural ecology of prehistoric parasitism on the Colorado Plateau as evidenced by coprology

The study of coprolites (desiccated feces) is recognized as a viable method for analyzing parasitism of prehistoric peoples. Eight species of helminth parasites, including nematodes, cestodes, and acanthcephalans, have been recovered from archaeological sites on the Colorado Plateau. The comparative analysis of parasitological findings illustrates the effects of changing subsistence strategies and varying life-style on prehistoric human parasitism. This comparative study is based on the analysis of coprolites recovered from one Archaic hunter-gatherer site and two Anasazi agricultural villages. Hunter-gatherers are represented by coprolites from Dust Devil Cave in south-eastern Utah. Coprolites of prehistoric agriculturalists from Antelope House in Canyon de Chelly, Arizona, and from Salmon Ruin in northwestern New Mexico were studied. The results demonstrate that helminth parasitism increased with the advent of agriculture. Between the agricultural sites, differences in patterns of excreta disposal, foraging behavior, and local ecology resulted in pronounced variations in both percentage of coprolites containing parasite remains and the number of parasite species represented.     

Trichuris sp. from 1,040 +/- 50-year-old Cervidae coprolites from the archaeological site Furna do Estrago, Pernambuco, Brazil

We present results of the paleoparasitological analysis of Cervidae coprolites that were recovered from the archaeological site Furna do Estrago, Pernambuco, Brazil. Trichuris sp. eggs were recovered from the coprolite samples dated 1,040 ± 50 years before present. This is the first record of Trichuris sp. in semiarid Cervidae, unexpectedly recorded in archaeological material.     

Insights from Characterizing Extinct Human Gut Microbiomes

In an effort to better understand the ancestral state of the human distal gut microbiome, we examine feces retrieved from archaeological contexts (coprolites). To accomplish this, we pyrosequenced the 16S rDNA V3 region from duplicate coprolite samples recovered from three archaeological sites, each representing a different depositional environment: Hinds Cave (∼8000 years B.P.) in the southern United States, Caserones (1600 years B.P.) in northern Chile, and Rio Zape in northern Mexico (1400 years B.P.). Clustering algorithms grouped samples from the same site. Phyletic representation was more similar within sites than between them. A Bayesian approach to source-tracking was used to compare the coprolite data to published data from known sources that include, soil, compost, human gut from rural African children, human gut, oral and skin from US cosmopolitan adults and non-human primate gut. The data from the Hinds Cave samples largely represented unknown sources. The Caserones samples, retrieved directly from natural mummies, matched compost in high proportion. A substantial and robust proportion of Rio Zape data was predicted to match the gut microbiome found in traditional rural communities, with more minor matches to other sources. One of the Rio Zape samples had taxonomic representation consistent with a child. To provide an idealized scenario for sample preservation, we also applied source tracking to previously published data for Ötzi the Iceman and a soldier frozen for 93 years on a glacier. Overall these studies reveal that human microbiome data has been preserved in some coprolites, and these preserved human microbiomes match more closely to those from the rural communities than to those from cosmopolitan communities. These results suggest that the modern cosmopolitan lifestyle resulted in a dramatic change to the human gut microbiome.     

南京汤山驼子洞鬣狗粪化石的孢粉分析

南京汤山驼子洞洞穴堆积中鬣狗粪化石含有相对丰富的孢粉,推测草食性哺乳动物食用带有孢粉的植物后,经鬣狗的猎杀、摄食、消化和排泄,孢粉较好地保存在鬣狗粪中,形成粪化石,成为洞穴地层的组成部分。虽然鬣狗粪化石中的孢粉含量不及湖沼相地层中丰富,但有几种植物的花粉保存较好且相对集中,如Pinus,Tsuga,Quercus,Carpinus,Artemisia,Cyperaceae,Polygonum,Polypodiaceae出现的频率较高。孢粉组合揭示早更新世南京地区植被主要是草原或森林-草地,气候凉干或半干旱半湿润,可以和地层中脊椎动物群反映的自然环境相互印证。在研究洞穴古环境的代用指标中,堆积物中的鬣狗粪化石中包含的孢粉是比较好的研究对象,对环境有一定的指示意义。     

Canid Coprolites from the Late Pleistocene of Hidalgo,Central Mexico: Importance for the Carnivore Record of North America

We present the first reported occurrence of canid coprolites from the late Pleistocene of central Mexico. The sample consists of five associated coprolites recovered from Quaternary deposits that crop out in southeastern Hidalgo, central Mexico. The material shows several of the typical features of canid feces, including elongated cylindrical shape, uncommon and nondistinct constrictions, and one tapered end. The coprolites are similar in size to feces of North American foxes, such as Vulpes macrotis, V. lagopus, V. vulpes, and Urocyon cinereoargenteus; likewise, their size might be comparable to those of the Pleistocene canid Canis cedazoensis. The content of the studied coprolites consists of numerous teeth and bone fragments referable to the pocket gophers Pappogeomys or Cratogeomys. The coprolite content is related to a mostly carnivorous diet comparable to that of C. cedazoensis, wolves (Canis lupus, C. rufus, and C. dirus), and some foxes (V. macrotis and V. velox). As a result, it is proposed that the potential trace maker of the coprolites could be a form similar to a carnivorous fox or a small dog. These trace fossils represent the first indirect evidence of canids in the late Pleistocene of Hidalgo and provide additional information regarding the geographic distribution of the Canidae in temperate areas of North America that now are part of central Mexico during the second half of the Pleistocene.     

Determination of prehistoric use of arrowhead (Sagittaria,Alismataceae) in the Great Basin of North America by scanning electron microscopy

Analyses of human coprolites from Dryden Cave, Nevada, indicated that the prehistoric population that inhabited the rockshelter included locally available herbs in their diet. Plant epidermal tissue remains have been identified as fragments from roots ofSagittaria (Alismataceae), a genus of semi-aquatic monocots. Other remains from the coprolites included fish bones, freshwater diatoms, seeds, and pollen. This assemblage supports the hypothesis that this prehistoric population principally exploited the available lacustrine resources of the area; comparisons with other archaeological sites suggests that this pattern had an extremely long history of development in the Great Basin.     

Molecular paleoparasitological diagnosis of Ascaris sp. from coprolites: new scenery of ascariasis in pre-Colombian South America times

Paleoparasitological studies using microscopy showed that Ascarisand Trichuris trichiura are the human intestinal parasites most found in archaeological sites. However, in pre-Columbian South American archaeological sites, Ascaris is rare. In this work we standardized a molecular methodology for Ascaris diagnosis directly from ancient DNA retrieved from coprolites. Using cythochrome b gene (142 bp) target, ancient DNA sequences were retrieved from South American samples, negative by microscopy. Moreover, the methodology applied was sensitive enough to detect ancient DNA extracted from 30 Ascaris eggs from an European coprolite. These results revealed a new scenery for the paleodistribution of Ascaris in South America.     

Vegetation reconstruction on the basis of pollen in Late Pleistocene hyena coprolites from San Teodoro Cave (Sicily, Italy)

Vegetation reconstruction based on pollen from coprolites of extinct spotted hyena (Crocuta crocuta spelaea, Goldfuss 1832) recovered from excavations carried out in 1998 at San Teodoro Cave (Sicily, Italy) supports previous indications of pre-Late Glacial conditions. Eight of the twelve coprolites analysed contained well-preserved pollen grains. There is a general similarity between the pollen contents from the coprolites but they show variability. They suggest a main vegetation type dominated by steppic taxa (Poaceae, Artemisia, Ephedra, Chenopodiaceae and Asteraceae) but also including arboreal taxa (Pinus and Cupressaceae). Low percentages of pollen of mesophilous woody taxa (Quercus, Betula, Abies, Alnus, Pistacia, among others) are noticeable, suggesting the existence of nearby refugia for temperate and Mediterranean vegetation. A reconstruction of the landscape, using the coprolite pollen record and other pollen records from Sicily and south Italy, shows the predominance, during the pre-Late Glacial, of a wooded steppe biome, with elements representing a variety of local environmental conditions.     

A palaeoparasitological analysis of rodent coprolites from the Cueva Huenul 1 archaeological site in Patagonia (Argentina)

The aim of the present study was to examine the parasite fauna present in rodent coprolites collected from Cueva Huenul 1 (CH1), northern Neuquén (Patagonia, Argentina), an archaeological site that provides stratified sequences of archaeological and palaeontological remains dating from the Late Pleistocene/Early Holocene Transition to the Late Holocene period. Twenty rodent coprolites collected from different sedimentary units from the site, with ages ranging from 13.844 ± 75-1.416 ± 37 years BP, were examined for parasites. Each coprolite was processed as a whole: rehydrated, homogenised, spontaneously sedimented and examined using light microscopy. The coprolites and the eggs of any parasites present were described, measured and photographed. In all, 158 parasite eggs were found in 10 coprolites. The faeces were positive for Viscachataenia quadrata Denegri, Dopchiz, Elissondo & Beveridge and Monoecocestus sp. Beddard (Cestoda: Anoplocephalidae) and for Heteroxynema (Cavioxyura) viscaciae Sutton & Hugot (Nematoda: Oxyuridae). The coprolites examined were tentatively attributed to Lagidium viscacia Molina (Mammalia, Rodentia, Caviomorpha, Chinchillidae). The life cycles of these parasites are discussed.     

Cryptosporidium,Giardia, and Cyclospora in ancient Peruvians

Twenty-two coprolites of human origin, collected from excavations along the north-central coast of Peru, were examined using fluorescent microscopy for the presence of fecal parasites, with emphasis on Cryptosporidium sp., Giardia sp., and Cyclospora sp. Three samples were positive. One coprolite dated between ca. 2,375 and 1,525 BC contained Giardia sp. cysts. This coprolite corresponded to the Peruvian preceramic period. Another positive coprolite ca. AD 770-830 corresponded to Epoch 3 of the Middle Horizon and contained Cryptosporidium sp. oocysts. The third positive coprolite (corresponding to the Middle Horizon. ca. AD 500-900) contained Giardia sp. cysts. This report demonstrates that Giardia sp. and Cryptosporidium sp. were present in Peruvian coastal populations for at least 4,300 and 1,100 BP.     

First record of Mesozoic scroll coprolites: classification,characteristics, elemental composition and probable producers

Multiple, small, cylindrical scroll coprolites having rounded and tapering ends and pertaining to a new ichnotaxon have been recovered from the Upper Triassic Tiki Formation of India. This is the first record of scroll coprolites from the Mesozoic. Based on cross‐sectional geometry, external surface textures, and internal morphology, these coprolites are subdivided into three morphotypes. The coprolites contain several kinds of undigested food material in the form of ganoid fish scales, teeth, lower jaw and skeletal remains of various osteichthyans, chondrichthyans, archosauriforms and indeterminate reptiles. These inclusions are embedded in the groundmass separated by thin mucosal layers. The groundmass contains abundant gas vesicles, and secondarily‐filled shrinkage cracks. EDS analysis shows that the overall composition of the coprolites reflects Ca, P, C and O, suggesting calcium phosphate mineralogy, though other elements such as Fe, Mn, Al, Si are present in lesser proportions. Based on their similarity with the scrolled faeces of extant euryhaline hammerhead sharks, it is deduced that these coprolites were produced by euryhaline hybodontid sharks. At least two hybodontid taxa, Lonchidion and Pristrisodus, show high prevalence in the Tiki vertebrate fauna, suggesting that these were the possible producers. As the coprolite inclusions contain remains of other aquatic animals, these carnivorous hybodonts constituted the dominant predators of the Tiki aquatic ecosystem.     

A molecular analysis of ground sloth diet through the last glaciation

DNA was extracted from five coprolites, excavated in Gypsum Cave, Nevada and radiocarbon dated to approximately 11 000, 20 000 and 28 500 years BP. All coprolites contained mitochondrial DNA sequences identical to a DNA sequence determined from a bone of the extinct ground sloth Nothrotheriops shastensis. A 157-bp fragment of the chloroplast gene for the large subunit of the ribulosebisphosphate carboxylase (rbcL) was amplified from the boluses and several hundred clones were sequenced. In addition, the same DNA fragment was sequenced from 99 plant species that occur in the vicinity of Gypsum Cave today. When these were compared to the DNA sequences in GenBank, 69 were correctly (two incorrectly) assigned to taxonomic orders. The plant sequences from the five coprolites as well as from one previously studied coprolite were compared to rbcL sequences in GenBank and the contemporary plant species. Thirteen families or orders of plants that formed part of the diet of the Shasta ground sloth could be identified, showing that the ground sloth was feeding on trees as well as herbs and grasses. The plants in the boluses further indicate that the climate 11 000 years BP was dryer than 20 000 and 28 500 years BP. However, the sloths seem to have visited water sources more frequently at 11 000 BP than at earlier times.     

Gar-Bitten Coprolite From South Carolina,USA

Coprolites can preserve a wide range of biogenic components. Of all the coprolites known from the fossil record, hitherto only two are known to preserve vertebrate tooth impressions (i.e., those of chondrichthyans). Here, a coprolite, from a thick lag deposit that includes a mixture of late Cretaceous, early Paleocene, and Plio-Pleistocene taxa at Clapp Creek in Kingstree, Williamsburg County, South Carolina, USA, preserves bite marks most consistent with having been made by a gar, Lepisosteus sp. (Lepisosteidae, Actinopterygii). This is the first-known coprolite to preserve actinopterygian tooth/bite marks. Aborted coprophagy seems unlikely; an accidental or serendipitous strike more likely describes the origin of the score marks over the surface of the coprolite. This coprolite also preserves small paired striations interpreted as evidence of coprophagy by an unknown organism.     

SL1 RNA gene recovery from Enterobius vermicularis ancient DNA in pre-Columbian human coprolites

Enterobius vermicularis, pinworm, is one of the most common helminths worldwide, infecting nearly a billion people at all socio-economic levels. In prehistoric populations the paleoparasitological findings show a pinworm homogeneous distribution among hunter-gatherers in North America, intensified with the advent of agriculture. This same increase also occurred in the transition from nomad hunter-gatherers to sedentary farmers in South America, although E. vermicularis infection encompasses only the ancient Andean peoples, with no record among the pre-Colombian populations in the South American lowlands. However, the outline of pinworm paleoepidemiology has been supported by microscopic finding of eggs recovered from coprolites. Since molecular techniques are precise and sensitive in detecting pathogen ancient DNA (aDNA), and also could provide insights into the parasite evolutionary history, in this work we have performed a molecular paleoparasitological study of E. vermicularis. aDNA was recovered and pinworm 5S rRNA spacer sequences were determined from pre-Columbian coprolites (4110 BC-AD 900) from four different North and South American archaeological sites. The sequence analysis confirmed E. vermicularis identity and revealed a similarity among ancient and modern sequences. Moreover, polymorphisms were identified at the relative positions 160, 173 and 180, in independent coprolite samples from Tulán, San Pedro de Atacama, Chile (1080-950 BC). We also verified the presence of peculiarities (Splicing leader (SL1) RNA sequence, spliced donor site, the Sm antigen biding site, and RNA secondary structure) which characterise the SL1 RNA gene. The analysis shows that the SL1 RNA gene of contemporary pinworms was present in pre-Columbian E. vermicularis by 6110 years ago. We were successful in detecting E. vermicularis aDNA even in coprolites without direct microscopic evidence of the eggs, improving the diagnosis of helminth infections in the past and further pinworm paleoepidemiological studies.     

The steroids of 2000-year-old human coprolites.

Six samples of human coprolites, some more than 2,000 years old, were analyzed for fecal steroid composition. Despite this very lengthy period of storage, the fecal steroids of coprolites were remarkably similar to those of stool samples collected today. The sterol nucleus was clearly rather stable under the dry environmental conditions of the Nevada Caves. The steroid content (microgram/g dried weight) of coprolite was low in comparison to that of modern man. The bile acid/cholesterol and plant sterol/cholesterol ratios of the coprolite, however, were similar to these ratios of the stools of modern man. In the six coprolites, an average 73% of the neutral steroids was digitonin-precipitable. This precipitate was composed of cholesterol and three plant sterols (campesterol, stigmasterol, and beta-sitosterol) and their bacteria-modified products. A portion of the neutral steroids had been converted to products tentatively identified as epimers of these steroids. Individual bile acids were identified in the coprolite. The bile acid composition of the coprolite was similar to that of the stool of modern man.     

High-resolution coproecology: using coprolites to reconstruct the habits and habitats of New Zealand's extinct upland moa (Megalapteryx didinus)

Knowledge about the diet and ecology of extinct herbivores has important implications for understanding the evolution of plant defence structures, establishing the influences of herbivory on past plant community structure and composition, and identifying pollination and seed dispersal syndromes. The flightless ratite moa (Aves: Dinornithiformes) were New Zealand's largest herbivores prior to their extinction soon after initial human settlement. Here we contribute to the knowledge of moa diet and ecology by reporting the results of a multidisciplinary study of 35 coprolites from a subalpine cave (Euphrates Cave) on the South Island of New Zealand. Ancient DNA analysis and radiocarbon dating revealed the coprolites were deposited by the extinct upland moa (Megalapteryx didinus), and span from at least 6,368±31 until 694±30 (14)C years BP; the approximate time of their extinction. Using pollen, plant macrofossil, and ancient DNA analyses, we identified at least 67 plant taxa from the coprolites, including the first evidence that moa fed on the nectar-rich flowers of New Zealand flax (Phormium) and tree fuchsia (Fuchsia excorticata). The plant assemblage from the coprolites reflects a highly-generalist feeding ecology for upland moa, including browsing and grazing across the full range of locally available habitats (spanning southern beech (Nothofagus) forest to tussock (Chionochloa) grassland). Intact seeds in the coprolites indicate that upland moa may have been important dispersal agents for several plant taxa. Plant taxa with putative anti-browse adaptations were also identified in the coprolites. Clusters of coprolites (based on pollen assemblages, moa haplotypes, and radiocarbon dates), probably reflect specimens deposited at the same time by individual birds, and reveal the necessity of suitably large sample sizes in coprolite studies to overcome potential biases in diet interpretation.     

The ichnogenous<Emphasis Type="Italic">Palaxius</Emphasis> (crustacean coprolite) and description of<Emphasis Type="Italic">P. hydranensis</Emphasis> n. sp. from the Upper Triassic (Norian part of “Pantokrator” limestone) of Hydra (Greece)

The fossil coprolite ichnogenusPalaxius is known with 22 species from the Permian up to the Tertiary. The diagnostic characteristics and the stratigraphie range of all species is given in this paper. The ichnogenusAgantaxia Kristan-Tollmann is a younger synonym of the ichnogenusPalaxius. The new crustacean coprolitePalaxius hydranensis n. sp. is described from the reef limestones of Norian part of “Pantokrator” limestone of the Island Hydra (Greece). This is the first report of coprolites from Hydra. The organism association of coprolite bearing limestone is mentioned.      

Paleocene Thalassinidea colonization in deep-sea environment and the coprolite Palaxius osaensis n. ichnosp. in Southern Costa Rica

Palaxius osaensis n. ichnosp., a new ichnospecies of crustacean coprolite is described. The coprolite is preserved in a 200-m thick Paleocene sequence in Southern Costa Rica that is largely dominated by pillow basalts. The studied sample is part of a seamount formed in the Pacific Ocean that was accreted to the Central American isthmus during the Eocene. The absence of lava vesicles, shallow-water deposits, and detrital sediments in the section suggest that the coprolites were deposited in a deep environment during the first stage of the development of the seamount. This represents one of the deepest occurrences of Thalassinidea coprolites reported in the literature and indicates that the producers of the coprolites, presumably some shrimps, developed the aptitude to colonize abyssal environments at least since Early Tertiary. The crustacean coprolites were encountered at a site which apparently lacked a food supply, although hydrothermal processes are believed to have provided the opportunity for a chemotrophic community to develop on the deepest part of the seamount. P. osaensis n. ichnosp. is also found in Colombia in late Cretaceous shallow-water sediments that notably contain Palaxius caucaensis coprolites (Micropaleontology 41 (1995) 85-88). Occurrences of P. osaensis n. ichnosp. deposited at both shallow and deep levels may possibly be related to an aptitude of some thalassinid organisms to have developed in various biotopes during the late Cretaceous-Paleocene.