The Dung File consists of a list of references dealing with pollen, parasites, and plant remains in coprolites and latrine fills from archaeological and palaeoenvironmental sites. The focus is on studies in North America.

The Dung File is subdivided into eleven sections: four depend on the origin of the deposits being investigated (Part 1: Mostly Human, Part 2: Mainly Mammal, Part 3: Animal Middens, Part 4: Other Critters), there are two broader categories, Part 5: General and Review Articles, Part 6: Field and Laboratory Methods, one focussed on theses, Part 7: Theses, and two focussed on modern comparative studies, Part 8: Comparative Studies - Human and Part 9: Comparative Studies - Mammal. Finally, there are a number of articles from news magazines and the popular press (Part 10: Popular Press and Commentary) and some less readily available items listed in Part 11: Conference Abstracts and Grey Literature.

The call numbers are for the library system at the University of Alberta. The remarks in black are my comments.

Part 2: Mainly Mammal

Agenbroad, L. D., and J. I. Mead 1989
Quaternary Geochronology and Distribution of Mammuthus on the Colorado Plateau. Geology 17:861-864.
AEU SCI QE 1 G3455 Area is basically the Four Corners region (Utah, Colorado, Arizona, New Mexico). Review of evidence to date. 41 localities with mammoth remains, 13 with dates (C14 and U series). Dates span around 28,000 - 10,000 yr BP. Includes mammoth dung samples dated from Bechan Cave. Latest dates average out at 11,270±65 yr BP. Possible that this marks the time of extinction. Dung analyses show that graminoids are major dietary components (grass, sedge, reed) with some woody shrubs. Reconstructed vegetation at Bechan Cave at this time is comparable to that in uplands to north and northwest (sagebrush steppe with riparian gallery forest). (18/05/2002).

Akeret, Ö., J. N. Haas, U. Leuzinger, and S. Jacomet 1999
Plant Macrofossils and Pollen in Goat/Sheep Faeces from the Neolithic Lakeshore Settlement Arbon Bleiche 3, Switzerland. The Holocene 9(2):175-182.
Site is near (formerly on) southwest shore of Lake Constance, NE Switzerland. Site was occupied for only 14 years (3384 - 3770 BC). Examined macroremains from 311 faeces, and pollen from 22 pellets. Rosaceae prickles (probably from Rubus) most common macroremains. Also silver fir (Abies alba) leaves and hazel (Corylus avellana) anthers. Samples also contain quantities of Abies and Corylus pollen. Few pellets contained parasite eggs. Macrofossil abundance is generally low. Data suggest winter as time of deposition. Two types of fodder consumed. Animals probably browsed on blackberry plants. Fir trees brought to village for construction and debranched; these branches then used for fodder. Animals foraging in area around village in daytime and brought back to the village for protection at night. Feed may have been supplemented by leaf fodder (e.g., in severe weather conditions). Authors comment that this "corresponds quite well with animal husbandry systems that were described from parts of the Alps at the beginning" of the 20th century (p. 180).

Akeret, Ö., and S. Jacomet 1997
Analysis of Plant Macrofossils in Goat/Sheep Faeces from the Neolithic Lake Shore Settlement of Horgen Scheller - An Indication of Prehistoric Transhumance? Vegetation History and Archaeobotany 6:235-239.
Site is on SW shore of Zürichsee, SE of Zürich, NE Switzerland. Three cultural phases identified between 3080 - 3030 BC. Report on upper layer. Examined 213 pieces from 10 samples. Few contained identifiable remains, suggesting winter deposition (i.e., lack of seeds). Rosaceae prickles are most abundant, probably from Rubus, also found fern sporangia. Blackberries retain green leaves all winter; some ferns do also. Animals allowed out to forage in clearings or hedgerows around settlement in day and brought back to the village at night. Authors suggest that lack of faeces indicating summer occupation may be related to transhumance. No evidence that animals were fed leaf fodder, despite abundance of twigs in cultural layers.

Akeret, Ö., and P. Rentzel 2001
Micromorphology and Plant Macrofossil Analysis of Cattle Dung from the Neolithic Lake Shore Settlement of Arbon Bleiche 3. Geoarchaeology 16(6):687-700.
Presents results of analyses of four cowpats recovered from the Arbon Bleiche 3 archaeological site. Examined pat micromorphologically through thin-sectioning. Contaminants (sand, hazel nut shells) became incorporated in pat along fissures though not part of cow's diet. Selected non- contaminated pieces for macrofossil analysis. Most of pats (>50%) is wood fragments (identifiable pieces are from Abies alba). No parasite eggs found. Micromorphological structure confirms that these objects are cowpats. Plant macros show that the diet was probably mostly woody plants, especially fir. Trees brought to the village for construction, when debranched, foliage fed to cattle. Other foods include mistletoe (probably mostly because this is parasitic on conifers), blackberry, and ferns. Probably represent autumn or winter consumption events (these fodders are not optimal for cattle). Cattle perhaps foddered rather than free-grazing because they need considerable nutrition to over-winter well. (01/04/2002).

Amorosi, T., P. C. Buckland, K. J. Edwards, I. Mainland, T. H. McGovern, J. P. Sadler, and P. Skidmore 1988
They Did Not Live by Grass Alone: The Politics and Palaeoecology of Animal Fodder in the North Atlantic Region. Environmental Archaeology 1:41-54.
Briefly mentions plant macroremains (presumably mainly from dung) as part of review of evidence for fodder types, especially in Norse settlements in Iceland and Greenland (in past millennium).

Araújo, Adauto, Luiz Fernando Ferreira, Ulisses Confalonieri, Marcia Chame and Benjamin Ribeiro 1989
Strongyloides ferreirai Rodrigues, Vincent & Gomes, 1985 (Nematoda, Rhabdiasoidea) in rodent coprolites (8,000-2,000 years BP), from archaeological sites from PiauÍ, Brazil. Memorias do Instituto Oswaldo Cruz 84(4):493-496.
Found eggs and larvae of Strongyloides ferreirai and eggs of Trichuris in coprolites from Rock Cavy (Kerodon rupestris), a large rodent. Coprolites were collected from six archaeological sites in São Raimondo Nonato, in northeast Brazil. Radiocarbon dates on sites span 2,000 to 8,000 yr BP. Identification of species that produced the coprolites was made by comparison with dung pellets collected from various modern rodents and small animals. Samples from all six sites yielded Strongyloides; three yielded Trichuris. Strongyloides larvae were identified after necropsy of modern Rock Cavy specimens; this has been described as a new species.

Bain, A. L., A. V. Morgan, J. A. Burns, and A. Morgan 1997
The Palaeoentomology of Rat's Nest Cave, Grotto Mountain, Alberta, Canada. In Studies in Quaternary Entomology - An Inordinate Fondness for Insects, edited by A. C. Ashworth, P. C. Buckland and J. P. Sadler, pp. 23-33. Quaternary Proceedings No. 5. Quaternary Research Association. John Wiley & Sons Ltd, Chichester, England, UK.
Insect remains were recovered from an organic accumulation spanning approximately last 2,000 years. Material has accumulated at the foot of a steep drop at rear of cave entrance (a natural trap). Deposit also yielded abundant faunal remains (34 vertebrate taxa). Packrats (bushy-tailed woodrat, Neotoma cinera) are present in the cave, but this is not strictly a midden. Insect fauna includes ants (Formicidae, 29%), flies (Muscidae, 29%), and beetles (Coleoptera, 63%). Of the beetles, 52% comprise Scarabaeidae (dung and chafer beetles), with 99% of these accounted for by one taxon, Aphodius congregatus. This is not indicative of any particular faunal species but is probably related to survival of some animals who fell into the pit and lingered for a while, plus availability of packrat dung. Most insects are not indicative of a cave environment but relate rather to the surrounding landscape. However, the beetle assemblage does include a few specimens of two cave-dwelling taxa.

Begler, E. B., and R. W. Keatinge 1979
Theoretical Goals and Methodological Realities: Problems in the Reconstruction of Prehistoric Subsistence Economies. World Archaeology 11(2):208-226.
Examines quantification of plant remains from two archaeological sites, through analysis of midden deposits. Sites are on north coast of Peru, dating to 1000 to 1534 AD, with an economy based on agriculture, supported by irrigation, and exploitation of marine resources. Sites are Cerro la Virgen and Medaños la Joyada. Found remains of 19 plant taxa. Also found coprolites from llama, guinea pig, and rodents (rat and/or mouse). No analysis of coprolites reported. Most of paper deals with issues of data analysis and quantification. (14/04/2007).

Bouchet, F., A. Araújo, S. Harter, S. M. Chaves, A. N. Duarte, J. L. Monnier, and L. F. Ferreira 2003
Toxocara canis (Werner, 1782) Eggs in the Pleistocene Site of Menez-Dregan, France (300,000-500,000) Years Before the Present. Memorias do Instituto Oswaldo Cruz 98 (Suppl. 1):137- 139.
Site is a collapsed cave in Brittany, northwest France. Examined sediment samples from five layers (5e, 7 and 9a) associated with human occupation. Found parasite eggs, Toxocara eggs, in sediment from layer 7. Size and morphology is consistent with Toxocara canis. May be from hyaena (Crocuta spelea) or another canid. Eggs well-preserved. One pollen grain (Carya) found in sediment samples. (01/06/2008).

Bouchet, F., C. Lefèvre, D. West, and D. Corbett 1999
First Paleoparasitological Analysis of a Midden in the Aleutian Islands (Alaska): Results and Limits. The Journal of Parasitology 85(2):369-372.
AEU SCI QL 757 J86 Buldir Island in western Aleutians. Examined organic sediments from two woodworking workshop sites and a house feature, dating between 13th and 17h centuries AD. Woodworking sites yielded parasite egg shells of either Diphyllobothrium or Schistocephalus (can't distinguish which). Diphyllobothrium associated with fish and also affects sea-lions (colony on the island). Schistocephalus would suggest presence of sea-birds (e.g., cormorants, gulls, and kittiwakes on the island). Also found nematode eggs, but could be free-living rather than from parasites. Hence, none of these can be definitively associated with humans. No parasite remains found in samples from the possible house structure. Plant remains (fungi, grass) also recovered from these samples, lending to support for archaeological interpretations of the features. (31/03/2002).

Braillard, L., M. Guélat, and P. Rentzel 2004
Effects of Bears on Rockshelter Sediments at Tanay Sur-les- Creux, Southwestern Switzerland. Geoarchaeology 19(4):343-367.
DOI: 10.1002/gea.10123 Many cave sediments in Alps have large amounts of phosphates. Origin of these is disputed: bat guano, cave bear dung, or bird guano. Some deposits were so rich they were mined. Studied cave is just south of southeast corner of Lac Léman. Cave fill is up to 80 cm thick and consists of four layers. Each layer sampled and examined for physical (including particle size, examination of granule morphometry) and chemical (including carbonates, humus, pH, phosphates) analyses and XRD. Also examined micromorphology of sediments from thin sections prepared from an intact monolith. Phosphate values are high and cannot be explained by presence of bone fragments in sediments. Sediments have multi-source origin, most are endokarstic, with some contribution from roof fall, some from alluvial sources. Layer 3 near base probably accumulated prior to last glacial maximum and sediments are disturbed and mixed by cave bears (Ursus spelaeus) digging lairs. Biogenic origin for phosphates, likely from bear dung and urine. Also found lynx [species not given, likely Felis lynx] coprolites. Did not find any cave bear coprolites. Bear carcasses may also have added to the phosphates in sediments. (09/06/2008).

Buckland, W. 1822
Account of an Assemblage of Fossil Teeth and Bones of Elephant, Rhinoceros, Hippopotamus, Bear, Tiger, and Hyaena, and Sixteen Other Animals; Discovered in a Cave at Kirkdale, Yorkshire, in the Year 1821: With a Comparative View of Five Similar Caverns in Various Parts of England, and Others on the Continent. Philosophical Transactions of the Royal Society of London 112:171-236.
A classic paper in geology. Buckland examined the faunal remains in this cave, noting that none of them showed signs of being abraded by water transport, and hence could not have been brought to the cave during the diluvial episode. This flood episode was the current explanation for transport of materials at the time; this is before acceptance of glaciation as a transport mechanism. Noted that the majority of the bones were broken, and some were encrusted with stalagtite material, but that none showed signs of being derived from the rock. He notes the remarkably good preservation of the bones. Identified 22 animal species, including carnivores, especially hyaena; estimates, based on teeth, that at least 75 individuals were represented. Notes that many of the broken bones have tooth marks, which match those of the hyaena. Concludes that the cave was probably a hyaena den. On p. 186-187, he describes finding hyaena dung, described as "solid calcareous excrement of an animal that had fed on bones", which he called "album graecum". These specimens are illustrated on Plate 24, Fig. 6. Identification confirmed by chemical tests. Buckland did not use the word "coprolite" here; he coined that word in a later paper (Buckland 1829). However, this is, if not the first, then one of the first descriptions of subfossil coprolites in geological context. Buckland also uses modern analogues, especially observations of the extant Cape Hyaena, to support his identification. Notes occurrance of bones of "elephant, rhinoceros, and hippopotamus", and speculates that hyaenas may have dragged parts of these carcases into the cave, because the opening was too small to admit them otherwise. Considers and rejects alternate explanations for the bone assemblage. Reviews evidence from other caves, both in Britain and in Europe, that have similar deposits. The paper contains superb illustrations, line drawings of the faunal remains. These were done by two ladies, identified as Miss Morland and Miss Duncombe. (25/11/2007).

Buckland, W. 1829
On the Discovery of Coprolites, or Fossil Faeces, in the Lias at Lyme Regis, and in Other Formations. Transactions of the Geological Society of London, Series II 3:223-236.
A classic paper in which Buckland coined the term "coprolites" to refer to fossil faeces. Most of paper discusses occurrence of coprolites with Ichthyosaur and Plesiosaur skeletons recovered from the Lias and presents the evidence on which he deduces their origin. Also describes probable food remains in coprolites, an early example of dietary analysis. Describes similar coprolites from other hard-rock formations and concludes by mentioning the remains from Kirkdale Cave, described in his earlier paper (Buckland 1822). Paper is illustrated by four very finely- drawn plates illustrating many specimens. (13/04/2009).

Burney, D. A., H. F. James, F. V. Grady, J. Rafamantanantsoa, Ramilisonina, H. T. Wright, and J. B. Cowart 1997
Environmental Change, Extinction and Human Activity: Evidence from Caves in NW Madagascar. Journal of Biogeography 24(6):755-767.
AEU SCI G 1 J855 Mostly an excavation report listing finds, primarily faunal reamins but also some archaeological materials, from two cave systems (Anjohibe and Anjohikely) in the northwest coastal region of Madagascar. Reports (p. 763) bones of an immature Archaeolemur (extinct form of lemur) associated with elliptical faecal pellets in cave Anjohikely 2. Attributed the faecal pellets to the lemur. Contents showed omnivorous diet, included bat bones (from Hipposideros commersoni), and plant remains including pollen and seeds. Pellets yielded a date of about 830 C14 yr BP. Note that this is a very late occurrence for this taxon. (24/05/2009).

Burney, D. A., G. S. Robinson, and L. P. Burney 2003
Sporormiella and the Late Holocene Extinctions in Madagascar. Proceedings of the National Academy of Sciences 100(19, September 16 2003):10800-10805.
Using Sporormiella spore abundance as a surrogate measure for megafaunal biomass. Looked at Sporormiella abundance in sediment cores throughout the island. Archaeological and other evidence shows human arrival around 2000 years ago, settlement began in semiarid SW and gradually spread into more mesic areas later. Pollen records track climate changes over last 35,000 yr BP. transition to dryer conditions began around 4000 yr BP, and change to more dry-adapted vegetation. Increase in burning also caused vegetation change, with increase in grasses and other open ground indicators. High values of Sporormiella in SW before human arrival. When humans arrive, Sporormiella values decline and disappear and charcoal increases. Decline in spores seem to happen before charcoal rise, by as much as perhaps two centuries. Sporormiella rises again after introduction of cattle in more recent centuries (dated differently in different areas). Few Sporormiella spores in sediments from interior highlands or NW rainforest through the late Holocene, suggesting that megafaunal biomass in these area was always lower than in SW. Some megafaunal were well-adapted to open environments. Megafauna concentrated in wooded savanna. Megafaunal biomass concentrated in wooded savanna. Megafaunal biomass low in uplands (ericoid heath) and lower land humid rainforest. (Not a taphonomic artifact of pollen preservation because Sporormiella abundance does increase in these areas after the introduction of cattle.) Human hunting pressure decreased faunal abundances, leading to increased fuel build-up and then fires. Areas transformed to thorny deserts and short grasslands, and in uplands to a depauperate steppe. Woodland only remaining in steep and rocky areas of the highland interior.

Buurman, J., B. van Geel, and G. B. A. van Reenen 1995
Palaeoecological Investigations of a Late Bronze Age Watering- place at Bovenkarspel, the Netherlands. In Neogene and Quaternary Geology of North-west Europe, edited by G. F. W. Herngreen and L. van der Valk. Meded. Rijks Geol. Dienst. 52.
Pit feature interpreted as a cattle watering place. (Appears analogous to a modern prairie dug-out). Macrobotanical and palynological investigations. Some remains derived from cattle dung. Carbonized remains (human activities) and uncarbonized. Results show mixed nature of feature in-fill (some redeposited older material). Also found eggs of parasite Trichuris (p. 267) and fungal spores from characteristic dung taxa (see pp. 265-266).

Camacho, C. N., J. S. Carrión, J. Navarro, M. Munuera, and A. R. Prieto 2000
An Experimental Approach to the Palynology of Cave Deposits. Journal of Quaternary Science 15(6):603-619.
AEU SCI QE 696 J86 Describes the palynology of surface sediments from two caves in SE Spain. Notes that dung has been an important sediment source in caves. Found parasite eggs, Trichuris, and also spores of Sordariaceae, Sporormiella. Fungi(Tilletia and Thecaphora) could also have been introduced in dung. Notes that some pollen contributed through bat guano. Most pollen types in the cave sediments can be correlated with regional vegetation (including agricultural/cultivated types such as Olea, olive). Dry floor sediments found to give best (in sense of representative) results. Greater degradation in samples from rear of cave. Speleothems, in this location, not found reliable for representative pollen assemblages. (04/07/2005).

Carrión, J. S., J. Brink, L. Scott, and J. Binneman 2000
Palynology and Palaeoenvironment of Pleistocene Hyaena Coprolites from an Open-air Site at Oyster Bay, Eastern Cape Coast, South Africa. South African Journal of Science 96:449-453.
Coprolites recovered from an "open-air" (= surface exposed?) site of Middle Stone Age, with archaeological materials and vertebrate faunal remains. Thought to be Last Glacial (near MIS 5a/4 boundary), about 70,000 yr BP. Site is within dunes and consists of palaeosol later revealed by wind erosion. Artifacts are consistent with Howieson's Poort substage of MSA. Hyaenas used the site soon after humans. Morphology and size of coprolites and identification of faunal remains suggest the coprolites are from brown hyaena (Hyaena brunnea). Fauna and pollen indicate three ecological types: open grassland, closed habitat, standing water habitat. Twelve coprolite samples analyzed for pollen, and spectra compared with one surface sample. Pollen concentrations are low; only six samples yielded reasonable numbers of grains. Thirty- two pollen and spore types were identified. Some contamination (e.g., by Pinus) probably by pollen washing into cracks in coprolites. Main pollen type is Myrica, with Poaceae and some pollen from Stoebe/Elytropappus. This latter type would appear to indicate cooler conditions than present. Time of lower sea level when site had a more "inland" character. (04/06/2006).

Carrión, J. S., J. A. Riquelme, C. Navarro, and M. Munuera 2001
Pollen in Hyaena Coprolites Reflects Late Glacial Landscape in Southern Spain. Palaeogeography, Palaeoclimatology, Palaeoecology 176:193- 205.
AEU SCI QE 500 P15 Spotted hyaena (Crocuta crocuta) coprolites from Las Ventanas Cave, southern Spain. AMS dating suggests coprolites date to the very late Pleistocene, just prior to Holocene transition. Twenty-five coprolites analyzed, only ten yielded enough pollen for reliable analysis. Others were either sterile or contained very few grains. Main pollen types are Pinus, Poaceae, Artemisia, with Juniperus and Lamiaceae. Variety of sources for pollen in droppings, probably mainly through stomach contents of large herbivores the hyaenas consumed. So the pollen reflects the prey's diet. Pollen assemblage is consistent with inferred Younger Dryas vegetation. (04/07/2005).

Carrión, J. S., L. Scott, A. Arribas, N. Fuentes, G. Gil-Romero, and E. Montoya 2007
Pleistocene Landscapes in Central Iberia Inferred from Pollen Analysis of Hyena Coprolites. Journal of Quaternary Science 22(2):191-202.
AEU SCI QE 696 J86 DOI: 10.1002/jqs.1024 Sampled hyaena coprolites from two sites in central Spain: Villacastín and Los Torrejones. Villacastín is a rockshelter which has yielded abundant faunal remains (35 taxa) including spotted hyaena (Crocuta crocuta ssp. intermedia) with an inferred date of Middle Pleistocene. Los Torrejones is a cave system that has yielded Homo cf. neanderthalenis remains and some Middle Palaeolithic lithics, also 19 other animal taxa including Crocuta crocuta spp. spelaea (cave hyaena), thought to date from Upper Pleistocene (but before last glaciation). Coprolites assigned to Crocuta on basis of shape, size and content (bone fragments). Analyzed 10 coprolite specimens from each site: 5 from Los Torrejones and 8 from Villacastín yielded pollen. Between 10 and 20 pollen taxa recovered. Pollen probably derived from prey animals and likely varies depending on how much of prey viscera were consumed. Pollen assemblages dominated by Poaceae and Pinus at both sites and evergreen oak (Quercus) at Villacastín. Assemblages suggest mid Pleistocene landscapes at Villacastín were patchy with grasslands and open forests with oaks (especially evergreen oak) and pines. Agrees with picture obtained from faunal remains. Upper Pleistocene vegetation inferred from Los Torrejones consists of steppe grasslands, open pine forests and parkland or savannah vegetation with pine. Agrees with pollen assemblages and vegetation reconstructions from other Upper Pleistocene sites in Spain, confirming age assignment. May suggest a refugium for oak during the start of the last glacial. Also shows a patchy landscape. Changing pollen spectra do not represent changing vegetation and a temporal sequence but sampling of contemporaneous patchy landscape, especially since hyaenas may range up to 50 km from den sites. (30/06/2008).

Carrión, J. S., L. Scott, T. Huffman, and C. Dreyer 2000
Pollen Analysis of Iron Age Cow Dung in Southern Africa. Vegetation History and Archaeobotany 9:239-249.
Heaps of slag-like material at Iron Age sites in southern Africa are the result of burning cow dung for fuel. Pollen preservation is generally poor in archaeological sites but good in the dung samples - burning appears to "seal in" the pollen contents. Compared dung from two areas, dated by C14 on other mater: Area 1 (1660 - 1810 AD) and Area 2 (350 - 1900 AD), with 9 modern dung samples for comparison. Area 1 pollen spectra: dominated by Poaceae, with Chenopodiaceae, and some Asteraceae and Cyperaceae. Little AP. Also spores, including Sordariaceae, and Trichuris microfossils. Samples from Area 2 were quite similar. Modern dung represented modern vegetation, including exotics (Eucalyptus, Cupressaceae, and Pinus), and also included Sordariaceae spores and Trichuris. Modern dung generally gave a good match for nearby modern vegetation. Assemblage shows a marked contrast with the Iron Age samples. Suggests that past vegetation was more open than present. Thought that high grass pollen content possibly reflecting good rainfall, in other words, a moisture rather than a temperature signal. Open areas around settlements more likely due to firewood gathering than overgrazing. Pollen spectra from more distant sites (springs) showed greater amounts of shrub and tree pollen. Suggests that area around kraals were open compared to more distant areas, which were more wooded but still more open than today. (24/07/2004).

Carrión, J. S., L. Scott, and E. Marais 2006
Environmental Implications of Pollen Spectra in Bat Droppings from Southeastern Spain and Potential for Palaeoenvironmental Reconstructions. Review of Palaeobotany and Palynology 140:175-186.
AEU SCI QE 901 R45 DOI: 10.1016/j.revpalbo.2006.03.007 Presents pollen assemblages from 34 bat guano samples from 9 caves in southeast Spain, plus 9 surface samples (1 from each locality). Nine bat species are the most likely cave inhabitants; all are insectivorous and up to 5 species may share a cave site. Hence pollen in guano likely from insects foraged by bats plus pollen from bats themselves deposited as part of grooming activity. Bats usually feed fairly locally around roost site (ranging up to 5 km) so pollen should give sampling of local vegetation. Inspection of pollen assemblages of guano and surface samples suggested each provided reasonable reflection of local vegetation. Differences between 9 sites reflect vegetation differences between those sites, which are in different ecozones. Pollen in bat guano samples is generally well-preserved (less than 10% indeterminable). Also shows higher taxonomic diversity and higher pollen concentrations than surface samples. Bat guano also includes pollen from zoophilous plant taxa that do not appear or are under-represented in the surface pollen assemblages. On the other hand, surface pollen samples may have higher representation of wind-pollinated taxa. Cluster analysis supported the conclusion that groupings are related to local vegetation. Groupings by taxa are less clear but may show ecological trends, e.g., forest taxa (humid) and xeric taxa, or temperature. Consider that bat guano may provide relatively reliable ecological snapshot of surroundings. (30/06/2008).

Charles, M. 1998
Fodder from Dung: The Recognition and Interpretation of Dung- Derived Plant Material from Archaeological Sites. Environmental Archaeology 1:111-122. Fodder: Archaeological, Historical and Ethnographic Studies, edited by M. Charles, P. Halstead, and G. Jones, vol. 1.
Bronze Age site of Abu Salabikh, southern Iraq (and observations on dung fuel use in modern village of same name), also data from Tell Brak. Attempts to assess contribution of dung (e.g., used as fuel) to the archaeobotanical assemblages. Sheep/dung pellets recognized in many archaeological samples from Abu Salabikh but few from Tell Brak. Tell Brak samples are consistent with derivation from crops and associated weeds, but Abu Salabikh samples contain large amounts of wild/weed seeds, possibly indicating derivation from dung. Formation processes of the assemblages were different at the two sites. Sheep/goats at Abu Salabikh may have been fed barley grain mixed with wheat chaff. May have implications for archaeological interpretation (i.e., growing crops for fodder rather than human consumption).

Charles, M., P. Halstead, and G. Jones (editors) 1998
Fodder: Archaeological, Historical and Ethnographic Studies. Environmental Archaeology 1. Oxbow Books, Oxford, England, UK 126 pp.
Studies are focused mainly on Europe and the Near East. A useful series of papers, many of which deal, wholly or partly, with dung (Amorosi et al., Smith, Haas et al., Karg, Anderson and Ertug-Yaras, Charles, Hall and Kenward).

Cremaschi, M., S. Di Lernia, and L. Trombino 1996
From Taming to Pastoralism in a Drying Environment: Site Formation Processes in the Shelters of the Tadrat Acacus Massif (Libya, Central Sahara). In Volume 3: Colloquium V - Numerical Analysis and Image Processing in Archaeobotany, Colloquium VI - Micromorphology of Deposits of Anthropogenic Origin, edited by L. Castelletti and M. Cremaschi, pp. 87-106. Published for International Union of Prehistoric and Protohistoric Sciences. A.B.A.C.O., Forli, Italy.
Surveyed wadis in the massif and recorded rockshelters or caves. Many of these contain well-preserved dung layers, dating to mid-late Holocene. Most of paper consists of stratigraphic and micromorphological descriptions of sediments in two caves: Uan Afuda and Uan Muhuggiag. Uan Afuda yielded sediments extending to early Holocene with a lower level in inner part of cave C14 dating to 8000 yr BP. Cave contained coprolites, and sediments also yielded spherulites. Dung is likely from ovicaprids (i.e., sheep or goat) but is not associated with pastoralism. Thought to be from wild animals, such as Barbary sheep (Ammotragus lervia). Sediments at Uan Muhuggiag date to middle Holocene. Here dung layers, both of cattle and sheep/goat, are likely related to pastoral activities. Sequence here shows a marked trend towards aridity in later Holocene. (09/10/2006).

Davis, O. K. 1987
Spores of the Dung Fungus Sporormiella: Increased Abundance in Historic Sediments and Before Pleistocene Megafaunal Extinction. Quaternary Research 28:290-294.
AEU SCI QE 696 Q24 Discusses the ecology and distribution of Sporormiella. Mostly found on herbivore dung. Became abundant after historic introduction of grazing animals in North America. Some heavily-grazed sites may not yield Sporormiella. Davis comments on the abundance of these fungal spores in late glacial sediments from six sites in the western US. Values suggest herbivore population densities similar to modern grazing densities. Associated vegetation is steppe or tundra. Also found in mammoth dung from Bechan Cave. Comments on coincidence of decline in abundance of Sporormiella spores with megafaunal extinction. Suggests spore abundance in marine cores (sediments derived partly from terrestrial sources) may provide an index for herbivore population densities.

Davis, O. K. 1987
Recent Developments in the Study of Arid Lands. Episodes 10(1):41-42.
AEU SCI QE 1 G3414 Discusses mammoth dung from Bechan Cave and mentions packrat midden analyses.

Davis, O. K., L. D. Agenbroad, P. S. Martin, and J. I. Mead 1984
The Pleistocene Dung Blanket of Bechan Cave, Utah. In Contributions in Quaternary Vertebrate Paleontology: A Volume in Memorial to John E. Guilday, edited by H. H. Genoways and M. R. Dawson, pp. 267-282. Carnegie Museum of Natural History Special Publication 8. Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA.
AEU SCI QE 741 C76 Cave discovered in 1982 by National Parks Service personnel. Site is in Glen Canyon area of SE Utah. Cave is on a steep sandstone slope and access to it is not easy. Extensive dung blanket, up to 40 cm thick, found with dung boluses that appeared to be analogous to African elephant (Loxodonta) droppings. Also found droppings from other herbivores and of ground sloth (recognizable by twig content. Appears to be similar to Cowboy Cave. Dung blanket underlain by sterile sand. No diagnostic large mammal bones found. Some small mammal bones found - cave is used by packrats. Cave had been used by Aboriginal people in the comparatively recent past - found corn cobs and metates but no pottery. No evidence of human occupation contemporaneous with dung blanket. Six radiocarbon dates from dung span about 11,600 -13,500 yr BP. Found hair and occasional hair masses during excavation - hair is consistent with mammoth, ground sloth, artiodactyls, horse and small mammals. Appears that main depositional interval was about a thousand years (11,700 - 12,900 years ago). Did pollen and macrofossil analysis on samples from excavation. Macros consist mainly of sedge and grasses, which form main component of dung samples. Macros representing riparian communities are present. Two sedge taxa (Carex lasiocarpa and Carex cf. lenticularis) occur further north (Idaho and Montana) at present. Also water birch (Betula occidentalis) now occurs only at higher elevations. Also found shrub macros (Sambucus [elderberry], Symphoricarpos [snowberry], Rosa [rose], Rubus [raspberry], and Ribes [currant]). Only small amount of spruce pollen but found cone galls caused by a gall aphid parasite (Chermes cooleyi). This is a particularly interesting finding - indirect evidence. Dune taxa include Opuntia polyacantha (cactus), Sclerocactus (Utah cactus), Atriplex canescens (salt bush), and Oryzopsis hymenoides (Indian rice grass). Abundant sage (Artemisia) pollen, up to 30%, in dung samples. Found macros of Artemisia cf. tridentata (big sage), which doesn't occur in the area today. Aquatic plant remains are more abundant towards the top of the dung layer. Quercus (oak) pollen is more abundant in top sample, whereas Picea (spruce) pollen is highest in lower samples and below the dung layer. Dung blanket indicates a time of vegetation transition. (04/06/2006).

Davis, O. K., J. I. Mead, P. S. Martin, and L. D. Agenbroad 1985
Riparian Plants were a Major Component of the Diet of Mammoths of Southern Utah. Current Research in the Pleistocene 2:81-82.
AEU HSS E 61 C97 Mammoth dung at Bechan Cave. Dung unit contains deposits from at least eight other species besides mammoth. Dates on layer indicate deposition between about 16,700-11,670 yr BP. Plant macroremains indicated mammoth dung consists primarily of a graminoid matrix. Saltbush, cactus and sagebrush remains in dung indicate xeric upland vegetation. Significant remains of aquatic/wetland plants (especially sedge and rush, with some naias, and horse-tail) indicate importance of riparian habitats.

Davis, O. K., and D. S. Shafer 2006
Sporormiella Fungal Spores, a Palynological Means of Detecting Herbivore Density. Palaeogeography, Palaeoclimatology, Palaeoecology 237:40- 50.
AEU SCI QE 500 P15 DOI: 10.1016/j.palaeo.2005.11.028 Two intervals when Sporormiella spores are in high abundance in lake sediments in western North America: late Pleistocene and in recent historical past. Sporormiella is a dung fungus and thus associated with grazing herbivores. It also has a distinctive spore. High percentages where grazers are corralled (e.g., 29% of pollen sum at Wildcat Lake, Washington). In grazed areas, but where animals have not been penned, usually about 3% - 4% of pollen sum. Can have very high values (up to 59%) in rock-shelter sites, which are used as bedding places by herbivores. Sometimes spores are absent in grazed areas, possibly due to local environmental factors. Generally, Sporormiella spores are either absent or in very low abundance in Holocene records. Reach above the 4% threshold value in sediments from many records of late Pleistocene age. Values match those associated with modern livestock grazing, suggesting megaherbivore population densities were about the same or similar to modern stocking densities. Spore decline appears to be associated with megafaunal extinction dates. (24/05/2008).

Eames, A. J. 1930
Report on Ground Sloth Coprolite from Doña Ana Co., New Mexico. The American Journal of Science 210:353-356.
AEU SCI QE 1 A51 Coprolite consists mainly of woody material (mostly stems, some roots) and some sand. Also found seeds, some hair, a few flower parts, and fern sporangia. Taxa identified include: Compositae (including achenes of Gutierrezia); Chenopodiaceae (including Atriplex fruits from three species); Malvaceae (including Sphaeralcea and perhaps Sida); and Cruciferae. Concludes that sloth was feeding on woody desert shrubs. Stems are young, indicating meal consumed in early season. Atriplex and Gutierrezia formed most of the meal. Results indicated vegetation similar to that found in area at present (but no evidence of desert succulents). Article is followed by a note from R. S. Lull conveying Eames' comments on preliminary analysis of dung of Neomylodon listai from Patagonia. Eames notes the presence of quantities of grass and sedge material, with no woody stems or roots.

Fonner, R. L. 1957
Appendix B, b: Mammal Feces from Danger Cave. In Danger Cave. Reprinted 1999, edited by J. D. Jennings, pp. 303. University of Utah Anthropological Papers Number 27. Society for American Archaeology No 14. University of Utah Press, Salt Lake City, Utah, USA.
AEU HSS E51 U88 No 27 Describes contents of four coprolite samples, thought to be from bear, though Fonner notes that they are morphologically indistinguishable from those in the previous report (Appendix B, c). All contain seeds of Allenrolfea occidentalis, with other material (some plant, some faunal). (31/07/2005).

Gill, Fiona L., Matthew P. Crump, Remmert Schouten and Ian D. Bull 2009.
Lipid analysis of a ground sloth coprolite. Quaternary Research 72:284-288.
AEU SCI QE 696 Q24 Took a sample from ground sloth (Nothrotheriops shastensis) dung specimen from Gypsum Cave. Age range likely between 11,000 - 36,000 RCYBP. Specimen contained yucca leaves. Faecal origin was confirmed by coprosterols in the extract. Lipid compound identified as epismilagenin was found in the extract. This compound is derived from saponins, probably from the vegetation the animal was consuming. Yucca and agave plants produce saponins that could have been the precursor source for this compound. The biomarker evidence thus supports the macrofossil evidence. So far this biomarker has only been found produced by modern ruminant animals. This therefore suggests characteristics of sloths' digestive systems which have hitherto not been known. (28/Feb/2013)

Graham, R. W., and J. I. Mead 1987
Environmental Fluctuations and Evolution of Mammalian Faunas During the Last Deglaciation in North America. In North America and Adjacent Oceans During the Last Deglaciation, edited by W. F. Ruddiman and H. E. Wright Jr, pp. 371-402. The Geology of North America Volume K-3. Geological Society of North America, Boulder, Colorado, USA.
This review concentrates mainly on the skeletal remains of mammals, but does mention dates on dung (including the mammoth dung studies) and faunal remains recovered from packrat middens. Many of the faunal assemblages discussed were retrieved from archaeological sites. Authors note that many Pleistocene assemblages are "non-analogue" groupings of taxa that are not found together today. The mammals, like the plants, are exhibiting an individualistic response to environmental change.

Haas, J. N., S. Karg, and P. Rasmussen 1998
Beech Leaves and Twigs used as Winter Fodder: Examples from Historic and Prehistoric Times. Environmental Archaeology 1:81-86.
A review of historical evidence for beech fodder (also hazel, birch, and alder). Early/Middle Bronze Age site of Fiavé-Carera (northern Italy). Sheep/goat dung recovered. Dendrochronological examination of twigs showed: young hazel twigs (1-3 years) collected in early spring; young beech twigs (4-12 years) also harvested in early growing season. Twigs with young leaves would be nutritious. Beech pollen not found in dung samples, so twigs harvested from young trees or from older trees that were already pollarded.

Hansen, R. M. 1978
Shasta Ground Sloth Food Habits, Rampart Cave, Arizona. Paleobiology 4(3):302-319.
AEU SCI QE 721 P145 Examined plant remains in dung from sloth (Nothrotheriops shastense) from Rampart Cave and some from Muav Caves. Notes that dung morphology is similar to those from other caves (especially Gypsum Cave) where dung found in association with sloth bones, as was also the case at Rampart Cave. Finite dates on deposits at Rampart Cave span about 36,000 to 10,700 C14 yr BP, with the lowermost level giving >40,000 C14 yr BP results. Two thick layers of sloth dung are separated by woodrat midden deposits, suggesting cave not used by sloths between about 23,000 to 13,000 yr BP. Not clear why this should be so. Also examined dung of modern animals in the region for comparison. Looked for plant epidermal fragments in dung samples and identified plants by comparison with modern reference material. Examined nutrient in samples of six of the most common plants in sloth dung to estimate dietary sufficiency. Examined 514 coprolites from Rampart Cave, and found 75 plant genera. Seven plants made up about 90% sloth diet: Sphaeralcea ambigua (Desert globemallow), Ephedra nevadensis (Nevada Mormon tea), Atriplex spp. (Saltbush), Acacia greggii (Catclaw acacia), Cactaceae, Phragmites communis (Common reed), and Yucca spp. All these plants are also found in the dung of modern herbivores in the area (bighorn sheep, cattle and burros (feral donkeys)). No change in sloth diets until the Late Pleistocene samples, which have much greater anounts of Ephedra. Samples from Muav Cave (not clear how many examined) were more mineralized and more difficult to analyze. Identified 20 plant genera. Most common were Common reed (Phragmites communis), Ash (Fraxinus), acacia, mesquite (Prosopis juliflora), Desert globemallow (Sphaeralcea ambigua), Nevada mormon tea (Ephedra nevadensis), and cactus. Modern plants show low digestible matter and high fibre content, generally not of high nutritional value, although digestible protein was likely nutritionally adequate, being highest for the Globemallow. Sloths apparently ate Globemallow, Mormon tea, and Saltbush all year and other plants only at specific times of year. Besides desert plants, sloths also ate plants typically found at seeps or shaded areas in region today. Caves were perhaps used as birthing sites. It was more likely that the sloths spent most of their time on the open range, since they were not well adapted to travel and would be unlikely to return to caves at end of day. They probably concentrated on browsing certain specific plants (those that are most common in their dung) and other plant material may have been largely ingested incidentally. (23/03/2008).

Hansen, R. M. 1980
Appendix III: Late Pleistocene Plant Fragments in the Dungs of Herbivores at Cowboy Cave. In Cowboy Cave, edited by J. D. Jennings, pp. 179-189. University of Utah Anthropological Papers Number 104. University of Utah Press, Salt Lake City, Utah, USA.
AEU HSS E51 U88 No 104 Looked at plant epidermal fragments from dung samples, also looked at seeds and hairs. Bison dung in level Ib comprised mainly grasses and sedges. Most identified hair was also from bison. Mammoth dung consisted almost entirely of grasses. Some possible horse dung also contained mainly grass. Ground sloth dung contained mainly Douglas fir needs and Serviceberry (Amelanchier) leaves. Some dung from a large herbivore remained unidentified (recorded as "camel-elk"). Some elk or deer dung contained mainly Serviceberry. Some hair of Bighorn Sheep was found but no firmly identified dung. Cottontail (Sylvilagus) pellets also found. Some canidae hairs and a few hairs tentatively identified as possibly human. (03/08/2003).

Hofreiter, M., J. L. Betancourt, A. P. Sbriller, V. Markgraf, and H. G. McDonald 2003
Phylogeny, Diet, and Habitat of an Extinct Ground Sloth from Cuchillo Curá, Neuquén Province, Southwest Argentina. Quaternary Research 59:364-378.
AEU SCI QE 696 Q24 DOI: 10.1016/S0033- 5894(03)00030-9 Dung from a new cave locality in far southwest Argentina. Thought to be from ground sloth on the basis of morphology. No skeletal remains found. Dung incorporated in a midden, cemented by urine, created by vizcachas (Lagidium sp., rodents in the chinchilla family). Plant DNA compared to GenBank sequences and those from a previous study (Hofreiter et al. 2000). Very little identifiable macro-plant material recovered from midden. Looked also for plant epidermal fragments and pollen (the latter apparently in one sample only). AMS dates from two labs differ by about a thousand years: 13,700 approx. vs. about 14,700 yr BP. Mitochondrial DNA results from dung were less clear than anticipated but overall matched most closely with sloth. Human and goat DNA also found and assumed to be later contaminants. Although grouped most closely with sloths, not clear which one. Dung is too small to be from known late Pleistocene types. Consider it is most likely from a "small nothrothere." This may be evidence of a hitherto unrecognized taxon of small sloths from South America, now extinct. Doesn't match any known skeletal material. Sloth dung yielded 13 plant DNA sequences, most in the Lamiales. Seven plant families identified. Plant cuticle analysis showed overlap with these results at the family level: Apiaceae, Asteraceae, Fabaceae, Lamiaceae, Anacardiaceeae, and Verbenaceae. Vizcaca pellets were dominated by four taxa (Naussavia, Poaceae, Mulinum, and Perezia) from the cuticle analysis. Midden matrix showed many of the same taxa: Mulinum, Perezia, Ephedra, Poaceae, and Junellia. Concludes that ground sloth was a browser not a grazer. Pollen in dung also yielded some taxa (e.g., Nothofagus) that may have resulted from long distance transport. No samples yielded evidence of Larrea, which is prevalent in the area today. Results compared well with data from another ground sloth side, Gruta del Indio, but are very different from Mylodon Cave. (01/06/2008).

Hofreiter, M., H. N. Poinar, W. G. Spaulding, K. Bauer, P. S. Martin, G. Possnert, and S. Pääbo 2000
A Molecular Analysis of Ground Sloth Diet Through the Last Glaciation. Molecular Ecology 9:1975-1984.
Analyzed 5 sloth coprolites from Gypsum Cave, Nevada, all of late Pleistocene age, and included data for another coprolite derived from a previous study (Poinar et al. 1998). Results presented for six coprolites. Also extracted DNA from sloth bone. DNA comparisons showed dung was from ground sloth. Two dung samples from about 28,500 C14 yr BP, two from around 20,000 C14 yr BP, and two from around 11,000 C14 yr BP. Used DNA sequences from 99 herbarium plant specimens from Gypsum Cave area for comparison with plant sequences extracted from dung. Also used data from database (GenBank) for comparison. Identified 13 families or orders in the plant DNA. Sequences show changes between the three time intervals in the preponderance of plant types. In oldest samples, Pinaceae, Moraceae, Capparales, and Poaceae are prominent. In middle samples, Capparales, Liliales, and some Poaceae. In youngest samples, Chenopodiaceae, Lamiales, Asteraceae, and Vitaceae. Suggests greater breadth of dietary resources used by sloths than previous macrofossil work had suggested. Also suggests vegetation (and climate) difference between the three intervals. Basically this study continues and extends work in previous study by Poinar et al. (1998). (01/06/2008).

Horrocks, Mark, Geoff Irwin, Martin Jones and Doug Sutton 2004
Starch grains of sweet potato (Ipomoea batatas) and bracken (Pteridium esculentum) in archaeological deposits from northern North Island, New Zealand. Journal of Archaeological Science 31:251-258.
AEU HSS CC 1 J86 Presents analysis of two starch crops (sweet potato [Ipomoea batatas] and bracken [Pteridium esculentum]) from soil in a stone mound and coprolites in archaeological context. Examined 13 samples. Five were coprolites: three from Great Barrier Island (outer Hauraki Gulf) which were from either human or dog, and two from Kohika (Bay of Plenty) which were probably from dog. Samples had previously been analysed for pollen and other microfossils. Mound soil samples from Pouerua site. Includes a detailed account of the starch extraction procedure. Starch granules in seven samples. Coprolites from Great Barrier Island contain starch grains and xylem (tissue) cells from sweet potato root. Starch from bracken rhizomes found in the Kohika coprolites. This confirms the cultivation of both crops in this area. Dating evidence from associated materials at the Great Barrier Island site suggests a post-450 RCYBP date. Starch in dog coprolites at Kohika probably from dogs consuming food scraps. Provides earliest evidence for use of bracken rhizome as a food source, likely around 1700 AD and later. Colour changes in starch granules suggest decomposition, perhaps due to cooking. (01/Mar/2013)

Horrocks, M., G. J. Irwin, M. S. McGlone, S. L. Nichol and L. J. Williams 2003
Pollen, Phytoliths and Diatoms in Prehistoric Coprolites from Kohika, Bay of Plenty, New Zealand. Journal of Archaeological Science 30:13-20.
AEU HSS CC 1 J86 Coprolites from a Maori site, near the coast, dating to late 17th century AD. Site is waterlogged (hence good material culture preservation) and on an island. Coprolites thought to be from dogs - analyses undertaken to confirm this. Fourteen coprolites (two of which further subsampled) analyzed. Pollen samples dominated by four taxa: Sonchus- type (puwha), Cyperaceae (sedge), Typha orientalis (raupo), and Pteridium esculentum (bracken) spores. Some samples also contained Coriaria (tutu) pollen. Phytolith assemblages are dominated by tree and grass-derived phytoliths. Diatom assemblages dominated by freshwater types. Large variety of pollen types and abundance of grass pollen adduced as evidence that coprolites deposited by dogs. High values of Sonchus-type pollen may indicate a dietary use of this plant (values are higher than expected from pollen analysis of core from adjacent wetland). A few seeds (Sonchus littoralis and Coriaria arborea) found in other Kohika coprolites, supporting dietary use of these plants. Conclude that coprolites were from dogs. (16/06/2006).

Hunt, C. O., G. Rushworth, D. D. Gilbertson, and D. J. Mattingly 2001
Romano-Libyan Dryland Animal Husbandry and Landscape: Pollen and Palynofacies Analyses of Coprolites from a Farm in the Wadi el- Amud, Tripolitania. Journal of Archaeological Science 28:351-363.
AEU PMC CC 1 J86 Examined coprolites from three intervals (Romano-Libyan, Arab, and modern). Farm is south of Lepcis Magna, in the interior, 200 km inland from the coast and on the desert margin. Modern inhabitants of the region are pastoralists. Archaeological remains suggest more intensive inhabitation in the past. Romano-Libyan farmsteads date mainly to 1st to 3rd centuries AD. Examined pollen in dung from sheep/goat, rodent, and one human sample. Older dung contained good pollen preservation but poor preservation in modern samples. Modern samples reflect modern aridland environment (dominated by Ephedra and Liliaceae), with abundant Lactuceae and some Poaceae. Late or post-Romano-Libyan rodent dung contains grass pollen and steppe indicators (e.g., Artemisia), plus pollen from crop plants (cereals and olives). Same pattern seen in coprolite of human origin. Sheep/goat dung of Romano-Libyan interval also shows pollen from steppe plants (notably Artemisia) and grass. Some contain large amounts of cereal pollen. Animals may possibly have been grazing on stubble or being fed straw. Suggests that animals were being confined or stalled and fed. Similar to modern animal husbandry in Mediterranean area. No evidence for vegetation or environmental change in this interval. Palynofacies assessment show abundance of degraded plant matter, as expected in dung. Most also showed fungal spores and hyphae, probably from fungi growing on dung after excretion. Also contain aquatic microfossils, probably from drinking water. Pollen from human coprolite is thermally mature (= darker) suggesting food cooking. (10/06/2006).

Jennings, J. D. 1980
Cowboy Cave. University of Utah Anthropological Papers Number 104. University of Utah Press, Salt Lake City, Utah, USA.
AEU HSS E 51 U88 Cowboy Cave, Utah, contained cultural material overlying a thick (50 cm) deposit of dung which included sloth, bison, mammoth, horse and a "camel-like ruminant." Cave situated in desert landscape of Colorado Plateau, with pinyon-juniper vegetation. Deposit consisted of five layers, each interspersed with a sterile sand layer. Dung layer dates to between 13 - 11 Ka BP. Data from four of the appendices described separately in this listing. These comprise: P. F. Hogan, Appendix IX: The Analysis of Human Coprolites from Cowboy Cave (pp. 201- 211), L. W. Lindsay, Appendix X: Pollen Analysis of Cowboy Cave Cultural Deposits (pp. 213-224), and Richard M. Hansen, Appendix III: Late Pleistocene Plant Fragments in the Dungs of Herbivores of Cowboy Cave (pp. 179-189), W. G. Spaulding and K. L. Petersen, Appendix II: late Pleistocene and Early Holocene Paleoecology of Cowboy Cave (pp. 163-177). (03/08/2003).

Jones, A. K. G., A. R. Hutchinson, and C. Nicholson 1988
The Worms of Roman Horses and Other Finds of Intestinal Parasite Eggs from Unpromising Deposits. Antiquity 62:275-276.
AEU HSS CC 1 A7 Reports evidence of Oxyuris equi (intestinal nematode of horses) "in Roman levels dated to c. AD 80-90 at the Annetwell Street site, Carlisle." Shows horses were on or near the site and may provide supporting evidence for identifying structures as stables. Also reports ova of human parasites (Trichuris and Ascaris) from Bronze Age material from Breen Down, Somerset, and from latrine pit fill (14th - 16th century AD) from Union Terrace, York (UK).

Jouy-Avantin, F., C. Combes, H. Lumley, J. C. Miskovsky, and H. Moné 1999
Helminth Eggs in Animal Coprolites from a Middle Pleistocene Site in Europe. The Journal of Parasitology 85(2):376-379.
AEU SCI QL 757 J86 Caune de l'Arago cave in the Pyrenees, records 300,000 years of occupation by humans and animals. One coprolite examined, dating earlier than 550,000 yr BP. Examined coprolite and attached sediment. More eggs in sediment than coprolite. Eggs of Dicrocoelidae, lower taxonomic identification not possible. Based on coprolite characteristics, the authors think it is from bear (faunal remains of both Ursus arctos and Ursus deningeri found in cave). (31/03/2002).

Karg, S. 1998
Winter- and Spring-foddering of Sheep/Goat in the Bronze Age Site of Fiavè-Carera, Northern Italy. Environmental Archaeology 1:87-94.
Lake-side habitation at time of occupation, some Neolithic in age, most is middle Bronze Age (1400 - 1300 B.C.). Twigs recovered from layers, also dung pellets examined for plant macroremains and pollen. Concludes that layers are animal dung and remains of foddering activity. Animals probably fed herbs/grasses, which were collected and dried for winter fodder, supplemented by leafy fodder from hazel and beech in the late winter/early spring.

Larkin, N. R., J. Alexander, and M. D. Lewis 2000
Using Experimental Studies of Recent Faecal Material to Examine Hyaena Coprolites from the West Runton Freshwater Bed, Norfolk, UK. Journal of Archaeological Science 27:19-31.
AEU HSS CC 1 J86 West Runton is the type site for the Cromerian Interglacial (early Middle Pleistocene). West Runton is just west of Cromer on the north coast of Norfolk, England. Site contains almost complete skeleton of Steppe Mammoth (Mammuthus trogontherii) associated with hyaena coprolites. Mammoth carcass probably deposited in slow-moving freshwater, and was scavenged by spotted hyaenas (Crocuta). Study designed to find out if coprolites were deposited in water or on dryland (i.e., did site dry out). Reviews studies on modern hyaena droppings (i.e., size, composition, density, etc.). Experimental set of 76 modern hyaena scats from Colchester Zoo. Compared chemical composition of fresh and ancient coprolites (14 specimens recovered from site), and coprolites of Crocuta crocuta from Boxgrove (another early Middle Pleistocene site). Fossil and modern coprolites show significant size difference, suggesting different populations. Various tests on modern droppings (e.g., soaking in water to see if they disintegrate) and flume experiments to see how transported in flowing water. Morphology and composition tend to support attribution to Crocuta crocuta. Larger sizes consistent with known Middle Pleistocene faunal remains from northern Europe (represents clinal variation in population). Experiments suggest that coprolites deposited in situ and not transported to the site by water. Also, could have been deposited in water because modern specimens don't disintegrate, even after prolonged immersion. So they were probably deposited by animals feeding on the carcass. (16/06/2006).

Laudermilk, J. D., and P. A. Munz 1934
Plants in the Dung of Nothrotherium from Gypsum Cave, Nevada. Carnegie Institution of Washington Publication 453:29-37. Contributions to Palaeontology Paper IV, vol. 453.
AEU SCI QE 746 C28 PP Dung balls associated with sloth remains from Gypsum Cave, recovered in 1930 and 1931. Previous research looked at two 100 cc (ml) samples of dung. About 80% was identified as Yucca. Reports on modern comparative collection of about 35 plant specimens gathered from vicinity of the cave. Also of plants from nearby in Clark Mountains at higher, moister elevation. Used these materials to undertake new examination of dung materials. Confirmed that most of fibre is from yucca and agave. Found large quantities of calcium oxalate crystals in finer fractions. Also recovered pollen from acetone washes. Compared plants recovered from sloth dung with those from burro (feral donkey) feeding in same area today. Obviously feed differently because yucca and agave were absent in burro dung. Sloth dung included Typha pollen showing that the animals ate wetland plants too. Yucca brevifolia and Agave utahensis were dominant in plant tissues from sloth dung. Neither plant grows at Gypsum Cave but both are found in Clark Mountains, about 3000 feet higher. Suggests cooler and moister environment when sloths were using Gypsum Cave. Dung was not dated (this is prior to availability of C14 dating). Paper includes detailed plant list with 14 plant taxa identified in sloth dung. Paper is followed by eleven plates, many showing plant epidermal cell structures. Note that publication date of volume is 1935, although this paper has a publication date on its title page of 1934. (24/07/2009).

Laudermilk, J. D., and P. A. Munz 1938
Plants in the Dung of Nothrotherium from Rampart and Muav Caves, Arizona. Carnegie Institution of Washington Publication 487:271-281. Contributions to Palaeontology Paper VII, vol. 487.
AEU SCI QE 841 C28 C Dung balls from newly- discovered Rampart Cave had a different plant assemblage to that recovered from Gypsum Cave samples. Caves (Rampart and Muav) partly excavated in 1937 by NPS (National Parks Service) staff. Dung blanket at Rampart Cave may be 6 feet or more thick. Little dung material at Muav Cave. Collected modern comparative plant specimens from vicinity of both caves. Yucca is not as abundant in these specimens, which consist mainly of Ephedra. One dung sample contained abundant grass and nematode worms and eggs, perhaps representing a pathological condition (i.e., a sick sloth). Muav specimens contained Yucca mohavensis; no Yucca in Rampart Cave specimens. Fine fractions contained hairs from Sphaeralcea and Atriplex. Also found spines and epidermal fragments of Opuntia, whereas none were found in Gypsum Cave material. Found Composite and grass pollen in acetone washes. Plant assemblage indicates an environment similar to present. Paper includes a list of plants found along trails to both caves (Muav and Rampart) and a list of plants found in the dung: 14 in Rampart and 12 in Muav specimens. Paper is followed 11 plates, most showing epidermal cell patterns. (24/07/2009).

Leroy, S. A. G., and M. J. Simms 2006
Iron Age to Medieval Entogamous Vegetation and Rhinolophus hipposideros Roost in South-Eastern Wales (UK). Palaeogeography, Palaeoclimatology, Palaeoecology 237:4- 18.
AEU SCI QE 500 P15 DOI: 10.1016/j.palaeo.2005.11.025 (Rhinolophus hipposideros = Lesser Horseshoe Bat). Bat guano incorporates pollen from three souces: pollen on insects eaten by bats, pollen from grooming, and airborne pollen blown into roost area. Bat species is endangered and found mainly in west UK, including Wales. Insect feeders, especially Diptera (midges) and don't usually forage more than 7 km from roost. Summer and winter roosts may be in different place; caves are often winter roosts. Samples from the Ogof Draenen cave system in Brecon Beacons National Park - a recent discovery (1994). Cave entrance blocked, perhaps by coal mining spoil; no longer used as a bat roost. Guano deposits are larger than any others known in UK. Cave system is connected to a small chamber, Siambre Ddu, that is still used as a summer roost for these bats. For comparison, collected two modern pollen samples (one moss polster, one lake mud), and 2 modern bat guano samples, from Siambre Ddu and from a roost at a nearby cave system, Agen Allwedd. Examined pollen from 10 samples of 19 collected from various parts of the cave system. Radiocarbon dates on samples span 2400 to 840 C14 yr BP. Fossil bat guano contains large amounts of AP, especially Hedera (Ivy) and Ilex (Holly). NAP percentages are comparatively low and are mainly Rosaceae types. Modern guano samples are also dominated by AP, but with more NAP types. Both Hedera and Ilex are insect-pollinated taxa and are rare in pollen assemblages from lake cores. Both are also rare in modern nearby vegetation. Fossil guano assemblages also contain higher proportions of oak (Quercus) pollen, suggesting more forested landscape in the past; present vegetation consists of more heathland and grassland. Assemblages also suggest that the cave was used as both a summer and winter roost. (27/12/2007).

Maher Jr, L. J. 2006
Environmental Information from Guano Palynology of Insectivorous Bats of the Central Part of the United States of America. Palaeogeography, Palaeoclimatology, Palaeoecology 237:19- 31.
AEU SCI QE 500 P15 DOI: 10.1016/j.palaeo.2005.11.026 Examined bat guano from Tumbling Creek Cave, southern Missouri. Inhabited by Grey Bat (Myotis grisescens), an insectivorous taxon, endangered. Bats forage up to 40 km from roost; surrounding vegetation is predominantly oak- hickory-pine woodland. Gives details of sampling and processing methods for bat guano. Centre of guano pile yielded date of about 2800 C14 yr BP. Some pollen likely trapped on bat fur; tested this by processing sample of bat fur for pollen. Experiment to see what pollen adheres to insects by trapping and processing night-flying moths for pollen; concludes they act as air-borne pollen traps. Same assumption made about insects on which bats feed. Processed sample of insect debris from a burnt-out light housing. Pollen assemblage compared well with upper sediment sample at Devils Lake. Thus bat guano contains representative sample of airborne pollen rain while they were feeding. If this is the case, then guano piles could contain a palaeoclimatic signal. Surface bat guano samples gives pollen assemblage similar to those from other sources (e.g., moss polsters, lake sediment surface samples). Bat guano deposition may not be continuous, this a guano pile may not be directly analogous to a lake core. Discusses possible techniques to explore stratigraphy of guano piles. [Note the different conclusions in this paper to others on bat guano. Does not report large proportion of entomophilous pollen. Possibly due to different species of bat and their feeding habits? If so, means that would need to know what species created fossil guano piles before their pollen spectra can be interpreted.] (27/12/2007).

Martin, P. S., B. E. Sabels, and D. Shutler Jr 1961
Rampart Cave Coprolite and Ecology of the Shasta Ground Sloth. The American Journal of Science 259:102-127.

Martin, P. S., R. S. Thompson, and A. Long 1985
Shasta Ground Sloth Extinction: A Test of the Blitzkrieg Model. In Environments and Extinctions: Man in Late Glacial North America, edited by J. I. Mead and D. J. Meltzer, pp. 5-14. Center for the Study of Early Man, Orono, Maine, USA.
AEU HSS QE 697 E62

Mead, J. I., L. D. Agenbroad, O. K. Davis, and P. S. Martin 1986
Dung of Mammuthus in the Arid Southwest, North America. Quaternary Research 25:121-127.
AEU SCI QE 696 Q24 Discusses dung of mammoth (Mammuthus) from Bechan Cave, southern Utah. Cave's name derived from Navajo word for "big feces." Dung layer, up to 40 cm thick, beneath 20 - 100 cm layer of loose sand and roof fall. Eight types of dung present; layer also yielded large amount of hair, including coarse hair identified as from mammoth. Recovered 17 complete or near-complete dung boluses produced by a very large mammal. Comparison with dung from modern elephants confirmed their identification as from mammoth. Dominated by coarse plant material, mainly (95%) grass and sedge. Some contained some woody material (saltbush [Atriplex sp.], sagebrush [Artemisia tridentata], water birch [Betula occidentalis], and blue spruce [Picea pungens]). Six C14 dates cluster between 12 - 13 Ka BP. Local riparian vegetation near cave included birch, elderberry (Sambucus sp.), wolfberry (Symphoricarpos sp.) and spruce. Community now found at higher elevations than cave. Regional vegetation (from pollen analysis) included big sagebrush, dwarf juniper (Juniperus communis), and oak (Quercus sp.). Suggests a sagebrush steppe. Conclude these data indicate a cooler and perhaps moister climate than present since modern equivalents are at more northerly latitudes or higher elevations.

Mead, J. I., L. D. Agenbroad, P. S. Martin, and O. K. Davis 1984
The Mammoth and Sloth Dung from Bechan Cave in Southern Utah. Current Research in the Pleistocene 1:79-80.
AEU HSS E 61 C97 Announcement of finding. Six C14 dates range from about 13,500 - 11,500 yr BP. Variety of other dung types reported besides mammoth.

Mead, J. I., L. D. Agenbroad, A. M. Phillips III, and L. T. Middleton 1987
Extinct Mountain Goat (Oreamnos harringtoni) in Southeastern Utah. Quaternary Research 27:323-331.
AEU SCI QE 696 Q24 Two metapodials and Oreamnos harringtoni dung on floor of rock shelter in Natural Bridges National Monument. Stratified deposits contain Oreamnos harringtoni dung and packrat middens. Dung referred to Oreamnos harringtoni on basis of size and morphology. Six layers (bottom to top) - Oreamnos harringtoni dung (with a "greater than" C14 date around 40 Ka) → packrat midden → Oreamnos harringtoni dung (dated around 23 Ka) → packrat midden (dated around 21 Ka) → mud → packrat midden (probably early Holocene, dated around 9.6 Ka). Pollen from both dung layers dominated by birch. Microhistological analysis indicated grass and sedges were ingested, but mainly bark, limber pine, and Douglas fir were dominant in the diet. Packrat middens include remains of conifers now found at higher elevations. All three layers contained extralocal taxa, but proportion is less in the most recent layer.

Mead, J. I., M. K. O'Rourke, and T. M. Foppe 1986
Dung and Diet of the Extinct Harrington's Mountain Goat (Oreamnos harringtoni). Journal of Mammalogy 67(2):284-293.
AEU SCI QL 700 J86 Describes the analysis of dung samples from two caves in Grand Canyon: Tse'an Bida Cave abd Tse'an Kaetan Cave. Compares dung morphology, size and weight to that of other grazing animals to establish its origin as likely from the extinct goat species. Defined dung pellet characteristics for this taxon as width:length ratio of >0.9, cuboid to subrectangular in shape, and >0.5 g weight. Also confirmed by association with faunal remains of the goat. Examined pellets for pollen and plant fragments (microhistological analysis) to estimate diets. Compared results of analyses with those from sediment samples from cave excavations to confirm consumption rather than ambient input. At Tse'an Bida Cave, pollen from dung is dominated by Compositae, Gramineae, and cf. Leptodactylon pollen. Gramineae and Compositae pollen is also abundant in cave sediment samples. Other pollen types in dung suggest consumption of vegetation in early spring to early summer. No pollen results included from Tse'an Keatan Cave. Microhistological analysis suggests grasses were major food item consumed at both sites. At Tse'an Bida Cave, these were Sporobolus, Festuca, and Oryzopsis, with some Carex, likely from the nearby canyon bottomlands, with about 10% conifer remains. At Tse'an Kaetan Cave, grass is again dominant, mainly Festuca and Agropyron, but conifers, notably Pinus, comprise about 25% of plant remains identified. Direct C14 dates on dung pellets span 11,800 to 30,600 yr BP, but most are from towards the younger end of that interval. Note that these dates help constrain time of extinction of the goat. Propose that the terminal date would be about 11,160 C14 yr BP. Concludes that goats lived mainly on xerophytic vegetation, grasses and other browse, which is different to the diet of related extant species. Diet does not give clues as to causes of extinction. (24/05/2009).

Navarro, C., J. S. Carrión, M. Munera, and A. R. Prieto 2001
Cave Surface Pollen and the Palynological Potential of Karstic Cave Sediments in Palaeoecology. Review of Palaeobotany and Palynology 117:245-265.
AEU SCI QE 901 R45 Examined sediments from six caves in southeast Spain. Samples from five yielded useful pollen spectra. Four of the caves contained dung and/or bat guano. Generally, pollen concentration is highest in sediments at cave mouth and declines to back of cave, because pollen is mostly from airborne sources. However, samples in cave interiors have greater amounts of likely animal-transported pollen types (often zoophilous taxa). Generally, pollen samples do seem to reflect the regional and local vegetation quite well, indicating that cave samples, even in karst areas, may provide useful representation of vegetation. Dry sediments had better pollen preservation than wet sediments, which have lower pollen concentrations and more degradation. (01/07/2007).

Nielsen, B. O., V. Mahler, and P. Rasmussen 2000
An Arthropod Assemblage and the Ecological Conditions in a Byre in the Neolithic Settlement of Weier, Switzerland. Journal of Archaeological Science 27:209-218.
AEU HSS CC 1 J86 DOI: 10.1006/jasc.1999.0448 At 5600 yr BP, this is the oldest byre in Europe. Settlement is in NE Switzerland, was on a small island in a shallow lake. Settlement at Weier was founded about 3800 BC. Animal husbandry was important, especially of cattle. Building 3 assumed to be a byre - contained layers of manure plus twig and leaf fragments. Dung layers separated by three wooden floors. Dung compressed to about 30 cm thick, probably thicker in the past. Examined 28 samples from 8 layers through the deposit. Found 54 taxa and 533 specimens of arthropods. Fly puparia, mites, and beetles were most abundant and formed about 98% of the assemblage. Majority of taxa and specimens found in middle layer of deposit. Puparia show that flies were actually breeding in the building. Mites have potential to yield more information with fine screening and better preservation. Adult beetles, especially flying rove beetles, may indicate environments outside the settlement, although few are definitive outdoor taxa. Only 37 beetle taxa recorded, which is low compared to other sites. Most are associated with decomposing organic matter. Few decomposer insects overall, perhaps indicating that byre was mucked out occasionally and manure spread on fields. Some of the taxa are diagnostic for stable manure. Most of fly types are ones that could be found in similar modern byres in the region. Also found liver fluke (Fasiola hepatica) in byre, confirming ruminants, cattle, present. Found remains of a biting louse (Damalinia bovis) associated with cattle. Usually most abundant in winter, so possibly byre used in winter. Large numbers of housefly (Musca domestica) puparia. Warm byres are ideal breeding locales for these flies, even in winter. Suggest byre was roofed not open, as a paddock. Appears the cattle fed leaf fodder. (24/11/2007).

Panagiotakopulu, E. 1999
An Examination of Biological Materials from Coprolites from XVIII Dynasty Amarna, Egypt. Journal of Archaeological Science 26(5):547-551.
AEU PMC CC 1 J86 Coprolites from Workmen's Village, dating to 14th century BC. Previous excavations found structures believed to be animal pens, with coprolites thought to be from pigs. Coprolites examined by other researchers who found parasite eggs of Ascaris and Taenia and plant remains, mostly from various grains. New examination of some coprolites from same site. Recovered plant remains of emmer (Triticum dicoccum), barley (Hordeum vulgare), and ryegrass (Lolium sp). Insects found included grain weevil (Sitophilus granarius) and the small-eyed flour beetle (Palorus ratzeburgi), both serious pests of stored grain. Concludes that animals fed on cereal waste and/or spoiled infested grain. Results support previous conclusion that penned animals were probably pigs. (20/04/2007).

Pirozynski, K. A., A. Carter, and R. G. Day 1984
Fungal Remains in Pleistocene Ground Squirrel Dung from Yukon Territory, Canada. Quaternary Research 22:375-382.
AEU SCI QE 696 Q24 Nest of arctic ground squirrel (Spermophilus parryii), found ca. 43 km se of Dawson City, and dated to 12,200±100 yr BP (GSC-2641). Dung (20 individual scats) yielded 8 fungal taxa, several of which are coprophilous.

Pirozynski, K. A., D. M. Jarzen, A. Carter, and R. G. Day 1988
Palynology and Mycology of Organic Clay Balls Accompanying Mastodon Bones - New Brunswick, Canada. Grana 27:123-139.
AEU SCI QK 658 G742 Hillsborough mastodon found in 1936 and dated around 37,000 yr BP. Attempts to determine the origin of clay balls found with remains. These have carbonate matrix with organic detritus containing wood fragments. Compared palynology and mycology of clay ball with that from a calcareous nodule found adhering to bones. Thirty palynomorph taxa identified. Pollen spectra of two samples are different. Clay ball dominated by Pinus, Picea and Cyperaceae pollen; nodule by Cyperaceae, then Pinus and Poaceae pollen. Sediments adjacent to faunal remains (peat and silt examined by Mott) yielded pollen spectra suggesting a boreal forest environment. Site where mastodon found probably a pond or small lake surrounded by an birch-alder swamp. Clay ball probably represents material ingested by the mastodon prior to death, though no convincing explanation is offered for the high amounts of mineral material (other than perhaps geophagy). Fungal spores are better represented and more diverse in the clay ball than the nodule and include types characteristic of the Sordariales (76% in clay ball, 41% in nodule), many types of which are associated with dung. Fungal spores include Gelasinospora, Neurospora, and Chaetomium. Other types recovered are characteristic of wood, coastal or brackish water, and marshland habitats. Authors note that preponderance of dung-type spores may be related to differential preservation. Paper contains photomicrographs of the fungal types recovered from the clay ball and nodule.

Poinar, H. N., M. Hofreiter, W. G. Spaulding, P. S. Martin, and B. A. Stankiewicz (and H. Bland, R. P. Evershed, G. Possnert, and S. Pääbo) 1998
Molecular Coproscopy: Dung and Diet of the Extinct Ground Sloth Nothrotheriops shastensis. Science 281(Number 5375, 17 July 1998):402-406.
AEU SCI Q 1 S41 Coprolite sample from Gypsum Cave, Nevada. AMS dated at 19,875±215 (Ua-11835). DNA analysis confirms origin as from a sloth, presumably N. shastensis. Further, DNA analysis indicates presence of at least 7 different plant types in the dung (their table, however, shows 8). These were identified at the Order level (Capparales, Cyperales, Liliales, Gentianales, Lamiales/Scrophulariales, Rhamnales, Malvales, Caryophyllales). Macrofossil analysis did not yield evidence of Capparales (the most dominant type according to the molecular analyses) or Gentianales, but did show Ephedra. Authors note that molecular analysis may reveal presence of plants that are difficult to detect through macroremains. Accompanied by a summary by Erik Stokstad in the News of the Week section (pp. 319- 320).

Rasmussen, P. 1989
Leaf-foddering of Livestock in the Neolithic: Archaeobotanical Evidence from Weier, Switzerland. Journal of Danish Archaeology 8:51-71.
Reviews ideas on leaf-foddering, tracing back to ideas about Neolithic elm decline and work of Troels-Smith. Notes that other arboreal pollen (AP) changes in Neolithic have also been ascribed to tree management (e.g., pollarding). Weier site (3100 - 2800 yr BC) in Switzerland provides strongest archaeological evidence for leaf-foddering including piles of twigs and byres with dung layers. Analyzed these dung layers to say what the animals were fed on. Dung samples contained parasite ova (Trichuris spp. and Fasciola hepatica). Liver fluke is evidence of ruminants. Thought dung layers are mainly from cattle. Dung layers contain large amounts of twigs. Eleven woody taxa are represented but proportions are consistent with preferred species where leaf-foddering is practised today, i.e., large preponderance (27%) of ash (Fraxinus), lime is also abundant (19%). Interestingly, elm is present in only very small amounts (3%). Ivy (Hedera) is also common and is known to be used for fodder today, especially in winter (it's an evergreen). Paper includes some pictures of modern experiments feeding cattle with leaf-fodder, providing residue very similar to that found at Weier. Explored whether age distribution of twigs could provide any indication of tree management system (e.g., pollarding or shredding). Most twigs (almost 70%) were 1-5 yrs old. Suggests selective harvesting and tree management. Cattle probably confined to byre in winter months. Elm wood was not used to any great extent in construction of the settlement either. But pollen diagram from nearby lake shows clear elm decline, increasing around 3100 yr BC. Decline cannot be attributed to timber use or fodder use. Concludes that elm decline was not caused by human activity. Can't conclusively prove that it was Dutch elm disease, either, although found one trunk with characteristic traces of galleries of elm bark beetle (Scotylus scotylus), the vector for the disease. (09/10/2006).

Rasmussen, P. 1993
Analysis of Goat/Sheep Faeces from Egolzwil 3, Switzerland: Evidence for Branch and Twig Foddering of Livestock in the Neolithic. Journal of Archaeological Science 20:479-502.
AEU PMC CC 1 J86 Site is on shore of former lake (Wauwilermoos) in northern Switzerland. Three radiocarbon dates span around 5600 - 5200 yr BP. Community practised arable agriculture, plus hunting, fishing, gathering, and also kept livestock (predominantly goat/sheep, but also pigs and cattle). Looked at pollen (15 samples) and plant macros (13 samples). Morphology of samples suggests all coprolites from goat rather than sheep. Pollen content (of 10 samples) dominated by Alnus and Corylus pollen. Two yielded whipworm ova. Rest, which is at most 20% of assemblage, is NAP, mostly open ground/disturbed land taxa. Other samples (5) also contained large amounts of Betula pollen and fern spores. For macros, most samples yielded wood fragments. (14/04/2002).

Robbins, E. I., P. S. Martin, and A. Long 1984
Paleoecology of Stanton's Cave, Grand Canyon, Arizona. In The Archaeology, Geology and Paleobiology of Stanton's Cave, Grand Canyon National Park, Arizona, edited by R. C. Euler, pp. 116-130. Chapter 11 in Grand Canyon Natural History Association Monograph No 6.
AEU HSS E 78 A7 A668 Deposits span last 18,000 years. Large faecal pellets attributed to Oreamnos harringtoni (Harrington's mountain goat). Change in size of faecal pellets at around 10,800 yr BP correlated with extinction of this taxon. Mountain sheep (Ovis canadensis) frequented the cave in Holocene. Plant remains in faecal pellets indicated that animals were eating xeric (dryland) plants, similar to types still found in region, throughout interval represented by the deposit. Pollen from cave earth provided more information on vegetation (and hence climate) change over this interval than did the faecal pellets.

Robinson, D., and B. Aaby 1994
Pollen and Plant Macrofossil Analyses from the Gedesby Ship - A Medieval Shipwreck from Falster, Denmark. Vegetation History and Archaeobotany 3:167-182.
Wreck dated to mid to late 13th century and is about 80% intact. Hull lined with wattle on which was up to 40 cm thick layer of organic material, apparently, from the smell, dung. Examined macros and pollen in three samples from this later. Plant macros generally well preserved. Identified between 26 and 82 taxa. Two samples were mostly bran, cereal remains, and seeds from weeds from agricultural and ruderal lands (between about 30% and 40% of assemblage). Some taxa from shore plants, pasture and meadows, and plants typical of roadsides and field boundaries (e.g., Lathyrus pratensis [Meadow vetchling]). Third sample was mostly wetland moss remains (predominantly Rhytidiadelphus squarrosus) and macros from wildland plants. Pollen (only two samples reported) concentration generally high and pollen well- preserved. 56 taxa identified, mostly from NAP, especially pollen from cultivated plants (around 10%), particularly Avena-type [oats]. Found a few ova (3) of Trichuris. Tree pollen is present in both samples (ca. 4% and 12%) whereas no tree macros found. Pollen probably reflecting regional vegetation, whereas macros give better representation of habitats where animals grazed. Results are consistent with identification of organic layer as dung. Ecology and distribution of plants represented gives probable area of origin as southern Baltic. Documentary records describe export of cattle from southern Scandinavia to Germany at this time, perhaps the ship was part of this trade. (27/04/2007).

Robinson, D., and P. Rasmussen 1989
Botanical Investigations at the Neolithic Lake Village at Weier, North East Switzerland: Leaf Hay and Cereals as Animal Fodder. In The Beginnings of Agriculture, edited by A. Milles, D. Williams and N. Gardener, pp. 149-163. Symposia of the Association for Environmental Archaeology No. 8. BAR International Series 496. BAR, Oxford, England, UK.
Midden material recovered from byres yielded house fly puparia (Musca domestica) and ova from Trichuris (whipworm) and Fasciola hepatica (liver fluke). Occupation around 3000 yr BC. Deposits around village and in transect upslope to nearby cultivated terrace yielded cereal remains, and weed seeds, and seeds and fruits of wild food plants. Present study re-examined byre materials and also layers from the transect. Confirmed probable faecal nature of byre materials. Did some experimental work with modern animals (goat, sheep, cow) to see what kind of feed might produce remains similar to those found in byre. Cows did not digest whole grain which passed through undamaged. Sheep and goat chewed grains and residues seemed more similar to those in byre. Twigs from byre manure are from ash (Fraxinus), lime (Tilia), and willow (Salix). Most twigs from 1 - 4 years old. Selective gathering of leafy fodder to ensure palatability. Probably fed cereal (perhaps pounded) as a supplement to leaf hay. (11/05/2002).

Scott, L. 1987
Pollen Analysis of Hyena Coprolites and Sediments from Equus Cave, Taung, Southern Kalahari (South Africa). Quaternary Research 28:144-156.
AEU SCI QE 696 Q24 Coprolites attributed to brown hyaena (Hyaena brunnea) by comparison with modern specimens. Four levels; lower three are older than about 7.5 Ka. Generally poor pollen preservation, especially in lower levels. Grass pollen abundant throughout. Both coprolites and sediment samples show greater amounts of AP towards upper parts of deposit. In coprolites, pollen probably represents incidentally ingested regional pollen although some probably from plant material (which hyaenas are known to consume). Pollen spectra suggest climate was cooler and wetter when sediments in lower levels were deposited with open shrubby grassland vegetation. Midsection deposited under warmer drier climate with shrubby vegetation and acacia savanna, upper level (1A) represents a thornveld vegetation under a climate similar to present.

Scott, L., and J. S. Brink 1992
Quaternary Palaeoenvironments of Pans in Central South Africa: Palynological and Palaeoecological Evidence. South African Geographer 19(1/2):22-34.
Pans are not defined but from the discussion appear to be closed basins, often wet, fed by springs or groundwater, often with saline deposits, and often subject to deflation. Pans often yield faunal and archaeological remains. A wide-ranging review but mentions coprolites from two sites: Deelpan and Florisbad. Four coprolites from spotted hyaena (Crocuta crocuta) at Florisbad, are probably of Late Pleistocene age. Pollen assemblages are dominated by grass pollen, suggesting open grassland. Coprolites of brown hyaena (Hyaena brunnea) recovered at Deelpan. Again the assemblages are dominated by grass pollen, suggesting open grassland. Dating difficult but likely late Holocene. Other faunal remains suggest permanent water available at that time. (17/06/2006).

Simons, E. L., and H. L. Alexander Jr 1964
The Age of the Shasta Ground Sloth from Aden Crater, New Mexico. American Antiquity 29(3):390-391.
AEU PMC CC 1 A6 Reports two C14 dates, one on desiccated tissue and the other on a coprolite specimen from the Ground Sloth (Nothrotherium shastense). Tissue date is 1000 C14 years younger than the coprolite date. Result attributed to contamination with alcohol, probably used as a preservative when the specimen was originally found. A previously-obtained young date on this tissue had led to speculaion about possible late survival of this species, beyond the extinction age for other megafauna. Coprolite C14 date was consistent with those from other megafauna, at about 11,000 yr BP. Specimen and coprolite found in a narrow shaft with no other faunal remains and hence assumed to be associated, from the same animal. (09/11/2008).

Smith, B. D. 1997
The Initial Domestication of Cucurbita pepo in the Americas 10,000 Years Ago. Science 276(Number 5314, 9 May 1997):932-934.
AEU SCI Q 1 S41 Evidence for domestication of squash (Cucurbito pepo) comes from five caves excavated in Mexico. Paper reports AMS dates on cucurbit remains from Guilá Naquitz Cave. Cucurbit remains from this cave show change in rind thickness, colour and peduncle size that indicate domestication. Change dated to 7000 C14 yr BP. Size of seeds suggests domesticated squash present by around 9000 C14 yr BP. Seed sizes compared to seed assemblage recovered from mastodon dung which is taken to represent wild squash. This change in seed size occurs earlier than the other morphological changes. Shows that different lines of evidence for domestication may lead to different conclusions. (03/08/2003).

Spaulding, W. G., and P. S. Martin 1979
Ground Sloth Dung of the Guadalupe Mountains. In Biological Investigations in the Guadalupe Mountains National Park, Texas, edited by H. H. Genoways and R. J. Baker, pp. 259-269. National Park Service, Proceedings and Transactions Series, No. 4. National Park Service, Washington, D.C., USA.

Spaulding, W. G., and K. L. Petersen 1980
Appendix II Late Pleistocene and Early Holocene Paleoecology of Cowboy Cave. In Cowboy Cave, edited by J. D. Jennings, pp. 163-177. University Utah Anthropology Paper Number 104. University of Utah Press, Salt Lake City, Utah, USA.
AEU HSS E 51 U88 No 104 Contains a detailed description of the stratigraphy of the basal dung layer. 32 pollen taxa identified from main dung layer (Ib), dominated by pollen of Gramineae and Cyperaceae, reflecting diet of herbivores. Contribution of pollen from non-dung sources (e.g., arboreal pollen) increases towards top of Stratum Ib and into overlying IIa which contains 29 taxa, including Picea and Douglas fir pollen. Also Populus pollen which is well preserved (probably because of dry cave environment). Top of Stratum IIa shows increasing abundance of Artemisia pollen and other xerophytic taxa. Macrofossil analysis found 15 taxa in Ib include aquatics, sedges, grasses, sagebrush, cactus, spruce, Douglas fir needles, and juniper twigs (Juniperus scopulorum). 27 taxa in macros from IIa including gambel oak leaves (Quercus gambellii), a tree which is common near the cave today. Radiocarbon dates showed that Ib spanned about 2000 yrs (i.e., 13 - 11 ka BP). Concludes that montane-type plant community existed around site in Late Pleistocene. The canyon bottom may have supported a perennial stream (wetland taxa and sedges). Community persisted until around 8700 yr BP. Large pollen concentrations suggest that the dung reflects food ingested during the growing (= flowering) season. (03/08/2003).

Spaulding, W. G., and T. R. Van Devender 1980
Appendix I Late Pleistocene Montane Conifers in Southeastern Utah. In Cowboy Cave, edited by J. D. Jennings, pp. 159 - 161. University of Utah Anthropological Papers Number 104. University of Utah Press, Salt Lake City, Utah, USA.
AEU HSS E51 U88 No 104 Dung yielded needles of spruce (Picea engelmannii or pungens) and Douglas fir (Pseudotsuga menziesii). Nearest modern representatives are km away and at higher elevations. Lowest elevation record of such taxa so far. (03/08/2003).

Sperry, C. C. 1957
Mammal Feces. In Danger Cave. Reprinted 1999, edited by J. D. Jennings, pp. 302. University Utah Anthropology Paper Number 27. Memoirs of the Society of American Archaeology No 14. University of Utah Press, Salt Lake City, Utah, USA.
AEU HSS E51 U88 No 27 Describes contents of 12 coprolite samples. Most identified as probably from coyote, others possibly from bear, bobcat, or wolf. and one sample of sheep dung. Many of the samples contain fibres of alkali bulrush (Scirpus paludosus) and pickleweed (Allenrolfea occidentalis). Deer and rabbit remains present in many of the carnivore coprolites. Sheep dung mainly wood fragments. (31/07/2005).

Taylor, E. L. 1955
Parasitic Helminths in Medieval Remains. The Veterinary Record 67:216-218 .
AEU SCI SF 601 V588 Material recovered during excavation in Winchester of a medieval tank or pit. Retrieved eggs of Trichuris trichiura, Ascaris lumbricoides, and Dicrocoelium dendriticum. Eggs are well preserved. Considers that the material is probably of animal origin, perhaps pigs. Ratio of Trichuris to Ascaris eggs is very high and opposite to expectations from modern samples from pigs; suggests this may be due to relative susceptibility to degradation. Alternatively, could be a latrine pit.

Thompson, R. S., T. R. Van Devender, P. S. Martin, T. Foppe, and A. Long 1980
Shasta Ground Sloth (Nothrotheriops shastensis Hoffstetter) at Shelter Cave, New Mexico: Environment, Diet, and Extinction. Quaternary Research 14:360-376.
AEU SCI QE 696 Q24 Lists known finds of ground sloth dung (Gypsum Cave, Nevada; Rampart and Muav Caves, Arizona; Aden Crater, New Mexico; High Sloth Caves and Williams Cave, Guadalupe Mountains, west Texas). Sloth dung at Shelter Cave dates between about 12,400 and 11,300 yr BP. Associated packrat midden samples indicate "a relatively xeric juniper woodland". Desert tortoise (Gopherus agassizi) remains in cave; dates show present at same time as ground sloth. This tortoise is not present in the area of the cave today. Plants in sloth dung inferred from cuticle analysis. Mormon tea (Ephedra) predominant in most samples. Other important plant remains include those of century plant (Agave) and Rosa-type. Pollen spectra from dung are quite different and include significant amounts of pollen from Juniperus, with Artemisia, Gramineae, and Compositae pollen. Pollen probably not related to deliberately consumed plants (i.e., not providing information about diet) but indicative of regional vegetation.

Valamoti, S. M. 2004
Plants and People in Late Neolithic and Early Bronze Age Northern Greece: An Archobotanical Investigation. British Archaeological Report (BAR) S1258. BAR (British Archological Reports), Oxford, England, UK 186 pp.
Report includes mention of seed assemblages typical of dung, possibly representing dung use as fuel, especially when charred. Includes dung-derived assemblages at Makri site and Mandalo, Makriyalos, and Arkadikos sites, archaeological sites in northern Greece. Discusses implications for investigating animal husbandry practices in Late Neolithic. (17/11/2007).

von den Driesch, A., and J. Boessneck 1983
A Roman Cat Skeleton from Quseir on the Red Sea Coast. Journal of Archaeological Science 10:205-211.
AEU PMC CC 1 J86 Most of paper consists of a discussion of the faunal remains and circumstances of burial. Roman period (1st - 2nd century AD). Cat was apparently very large for a house cat. Skeleton also associated with dung balls. Intestinal contents included remains of at least five rats (Rattus rattus), with remains of a sixth rat found in a dung ball. A very large meal for a cat! Provides indisputable evidence for presence of rat at this period. (10/06/2006).

Yll, R., J. S. Carrión, A. C. Marra, and L. Bonfiglio 2006
Vegetation Reconstructions on the Basis of Pollen in Late Pleistocene Hyena Coprolites from San Teodoro Cave (Sicily, Italy). Palaeogeography, Palaeoclimatology, Palaeoecology 237:32- 39.
AEU SCI QE 500 P15 DOI: 10.1016/j.palaeo.2005.11.027 Coprolites are from spotted hyena (Crocuta crocuta) on basis of morphology and included bone fragments. Also found spotted hyena faunal remains in cave. 12 coprolite samples from different levels examined for pollen; 8 yielded countable pollen spectra. Assemblages are dominated by AP (mainly Poaceae), AP is mainly Pinus and Cupressaceae. Pollen spectra agree with those from regional pollen records. [Note: No C14 dates, only identified as from "pre-Late Glacial" levels.] (27/12/2007).

Number of citations: 81

Mainly Mammal


This presentation has been compiled and is © 1998-2015 by
Alwynne B. Beaudoin (
Last updated July 17, 2015
You are visitor #8613

For optimal viewing, the following browsers are recommended:
Firefox, Google Chrome, Opera, Safari and SeaMonkey.