Literature on Coprolites

The Principal Literature on Coprolites, Other Bromalites, and Modern Feces

This list is still very incomplete (199 items). Check for updates from time to time. The list is divided into two chapters: ‘Bromalites‘ and ‘Modern feces‘.



Aceñolaza, F.G., 2012. Neogene coprolites (Upper Miocene–Lower Pliocene) of Entre Rios Province, northeastern Argentina. Sci. Interfluvius 3 (1), 41–53.

Ambwani, K., Dutta, D., 2005. Seed-like structure in dinosaurian coprolite of Lameta Formation (Upper Cretaceous) at Pisdura, Maharashtra, India. Curr. Sci. India 88, 352–354.

Anagnostakis, S., 2013. Upper Cretaceous coprolites from the Münster Basin (northwestern Germany) – a glimpse into the diet of extinct animals. Dissertations in Geology at Lund University, Msc. Thesis, no 357, p. 30.

Appelt, S., 2013. Paléomicrobiologie des coprolithes. Ph.D. Thesis, Aix-Marseille Université, Faculté de Médecine, p. 261.

Appelt, S., Armougom, F., Le Bailly, M., Robert, C., Drancourt, M., 2014. Polyphasic analysis of a Middle Ages coprolite microbiota, Belgium. PLoS ONE 9 (2), e88376. doi: 10.1371/journal.pone.0088376

Appelt, S., Fancello, L., Le Bailly, M., Raoult, D., Drancourt, M., Desnues, C., 2014. Viruses in a 14th-century coprolite. Appl. Environ. Microbiol. 80, 2648–2655. doi: 10.1128/AEM.03242-13


Badal, E., Atienza, V., 2005. Análisis microscópico de coprolitos de herbívoros hallados en contextos arqueológicos. VI Congreso Ibérico de Arqueometría, Avances en Arqueometría, 283–293.

Bajdek, P., 2013. Coprolite of a durophagous carnivore from the Upper Cretaceous Godula Beds, Outer Western Carpathians, Poland. Geol. Q. 57 (2), 361–364. doi: 10.7306/gq.1094

Bajdek, P., Owocki, K., Niedźwiedzki, G., 2014. Putative dicynodont coprolites from the Upper Triassic of Poland. Palaeogeogr. Palaeoclimatol. Palaeoecol. 411, 1–17. doi: 10.1016/j.palaeo.2014.06.013

Bajdek, P., Qvarnström, M., Owocki, K., Sulej, T., Sennikov, A.G., Golubev, V.K., Niedźwiedzki, G., 2016. Microbiota and food residues including possible evidence of pre-mammalian hair in Upper Permian coprolites from Russia. Lethaia 49, 455–477. doi: 10.1111/let.12156

Bajdek, P., Owocki, K., Sennikov, A.G., Golubev, V.K., Niedźwiedzki, G., 2017. Residues from the Upper Permian carnivore coprolites from Vyazniki in Russia – key questions in reconstruction of feeding habits. Palaeogeogr. Palaeoclimatol. Palaeoecol. 482, 70–82. doi: 10.1016/j.palaeo.2017.05.033

Baldanza, A., Bizzarri, R., Famiani, F., Monaco, P., Pellegrino, R., Sassi, P., 2013. Enigmatic, biogenically induced structures in Pleistocene marine deposits: a first record of fossil ambergris. Geology 41 (10), 1075–1078. doi: 10.1130/G34731.1

Brachaniec, T., Niedźwiedzki, R., Surmik, D., Krzykawski, T., Szopa, K., Gorzelak, P., Salamon, M.A., 2015. Coprolites of marine vertebrate predators from the Lower Triassic of southern Poland. Palaeogeogr. Palaeoclimatol. Palaeoecol. 435, 118–126. doi: 10.1016/j.palaeo.2015.06.005

Bravo-Cuevas, V.M., Morales-García, N.M., Barrón-Ortiz, C.R., Theodor, J.M., Cabral-Perdomo, M.A., 2017. Canid Coprolites from the Late Pleistocene of Hidalgo, Central Mexico: Importance for the Carnivore Record of North America. Ichnos, 1–11. doi: 10.1080/10420940.2016.1270209

Broughton, P.L., 1981. Casts of vertebrate internal organs from the Upper Cretaceous of western Canada. J. Geol. 89, 741–749.

Broughton, P.L., 2016. Enigmatic origin of massive Late Cretaceous-to-Neogene coprolite-like deposits in North America: a novel palaeobiological alternative to inorganic morphogenesis. Lethaia. doi: 10.1111/let.12186

Broughton, P.L., Simpson, F., Whitaker, S.H., 1978. Late Cretaceous coprolites from western Canada. Palaeontology 21 (2), 443–453.

Buckland, W., 1835. 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, second series 3, 223–232.

Burmeister, K.C., 2000. Paleogeographic and biostratigraphic implications of new early Mesozoic terrestrial vertebrate fossils from the Poamay site; central Morondava Basin, Madagascar. Msc. Thesis, p. 109, University of California, Santa Barbara, USA.


Calvo, J.O., 1994. Gastroliths in sauropod dinosaurs. Gaia, 10, 205–208.

Cantrell, A.K., Suazo, T.L., Spielmann, J.A., Lucas, S.G., 2012. Vertebrate coprolites from the Lower Permian (Middle Wolfcampian) Gallina Well locality, Joyita Hills, Socorro Country, New Mexico. NMMNH Bull. 57, 197–201.

Cano, R.J., Rivera-Perez, J., Toranzos, G.A., Santiago-Rodriguez, T.M., Narganes-Storde, Y.M., Chanlatte-Baik, L., García-Roldán, E., Bunkley-Williams, L., Massey, S.E., 2014. Paleomicrobiology: revealing fecal microbiomes of ancient indigenous cultures. PLoS ONE 9, e106833.

Cerda, I.A., 2008. Gastroliths in an ornithopod dinosaur. Acta Palaeontol. Pol. 53 (2), 351–355.

Chin, K., 1997. Coprolites, in: Currie, P.J., Padian, K. (Eds.), Encyclopedia of Dinosaurs. Academic Press, pp. 147–150.

Chin, K., 2002. Analyses of coprolites produced by carnivorous vertebrates, in: Kowalewski, M., Kelley, P.H. (Eds.), The Fossil Record of Predation. Paleontological Society Special Papers, 8, pp. 43–49.

Chin, K., 2007. The paleobiological implications of herbivorous dinosaur coprolites from the Upper Cretaceous Two Medicine Formation of Montana: why eat wood? Palaios 22, 554–566. doi: 10.2110/palo.2006.p06-087r

Chin, K., 2007. Thin section analysis of lithified coprolites (fossil feces). Microsc. Microanal. 13 (Suppl. 2), 504–505. doi: 10.1017/S143192760707540X

Chin, K., Bishop, J.R., 2007. Exploited twice: bored bone in a theropod coprolite from the Jurassic Morrison Formation of Utah, U.S.A. SEPM Spec. P. 88, 379–387.

Chin, K., Gill, B.D., 1996. Dinosaurs, dung beetles, and conifers: participants in a cretaceous food web. Palaios 11, 280–285.

Chin, K., Kirkland, J.I., 1998. Probable herbivore coprolites fromthe Upper Jurassic Mygatt-Moore Quarry, Western Colorado. Mod. Geol. 23, 249–275.

Chin, K., Tokaryk, T.T., Erickson, G.M., Calk, L.C., 1998. A king-sized theropod coprolite. Nature 393, 680–682.

Chin, K., Eberth, D.A., Schweitzer, M.H., Rando, T.A., Sloboda, W.J., Horner, J.R., 2003. Remarkable preservation of undigested muscle tissuewithin a Late Cretaceous tyrannosaurid coprolite from Alberta, Canada. Palaios 18 (3), 286–294.

Chin, K., Bloch, J., Sweet, A., Tweet, j., Eberle, J., Cumbaa, S., Witkowski, J., Harwood, D., 2008. Life in a temperate Polar sea: a unique taphonomic window on the structure of a Late Cretaceous Arctic marine ecosystem. P. Roy. Soc. B-Biol. Sci. 275, 2675–2685. doi: 10.1098/rspb.2008.0801

Chin, K., Hartman, J.H., Roth, B., 2009. Opportunistic exploitation of dinosaur dung: fossil snails in coprolites from the Upper Cretaceous Two Medicine Formation of Montana. Lethaia 42, 185–198.

Codorniú, L., Chiappe, L.M., Cid, F.D., 2013. First occurrence of stomach stones in pterosaurs. J. Vertebr. Paleontol. 33 (3), 647–654. doi: 10.1080/02724634.2013.731335


Dentzien-Dias, P.C., de Figueiredo, A.E.Q., Horn, B., Cisneros, J.C., Schultz, C.L., 2012. Paleobiology of a unique vertebrate coprolites concentration from Rio do Rasto Formation (Middle/Upper Permian), Paraná Basin, Brazil. J. S. Am. Earth Sci. doi: 10.1016/j.jsames.2012.09.008

Dentzien-Dias, P.C., Poinar, G.Jr., de Figueiredo, A.E.Q., Pacheco, A.C.L., Horn, B.L.D., Schultz, C.L., 2013. Tapeworm Eggs in a 270 Million-Year-Old Shark Coprolite. PLoS ONE 8 (1), e55007. doi: 10.1371/journal.pone.0055007

Dentzien-Dias, P., Poinar, G., Francischini, H., 2016. A new actinomycete from a Guadalupian vertebrate coprolite from Brazil. Historical Biology. doi: 10.1080/08912963.2016.1241247

Diedrich, C.G., 2012. Typology of Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) coprolite aggregate pellets from the European Late Pleistocene and their significance at dens and scavenging sites. NMMNH Bull. 57, 369–377.

Doyle, P., Macdonald, D.I.M., 1993. Belemnite battlefields. Lethaia 26, 65–80.

Duffin, C.J., 2012. The earliest published records of coprolites. NMMNH Bull. 57, 25–28.

Dutta, D., Ambwani, K., 2007. Capers: a food for upper Cretaceous dinosaurs of Pisdura, India. Curr. Sci. India 92, 897–899.


Edwards, D., Selden, P.A., Axe, L., 2012. Selective feeding in an Early Devonian terrestrial ecosystem. Palaios 27, 509–522. doi: 10.2110/palo.2011.p11-094r

Eriksson, M.E., Lindgren, J., Chin, K., Månsby, U., 2011. Coprolite morphotypes from the Upper Cretaceous of Sweden: novel views on an ancient ecosystem and implications for coprolite taphonomy. Lethaia 44, 455–468. doi: 10.1111/j.1502-3931.2010.00257.x


Farlow, J.O., Chin, K., Argast, A., Poppy, S., 2010. Coprolites from the Pipe Creek Sinkhole (Late Neogene, Grant Country, Indiana, USA). J. Vertebr. Paleontol. 30 (3), 959–969.

Fiorelli, L.E., Ezcurra, M.D., Hechenleitner, E.M., Argañaraz, E., Taborda, J.R.A., Trotteyn, M.J., Belén von Baczko, M., Desojo, J.B., 2013. The oldest known communal latrines provide evidence of gregarism in Triassic megaherbivores. Scientific Reports 3, 3348. doi: 10.1038/srep03348

Fisher, D.C., 1981. Crocodilian scatology, microvertebrate concentrations, and enamel-less teeth. Paleobiology 7 (2), 262–275.

Fisher, D.C., 1981. Taphonomic interpretation of enamel-less teeth in the Shotgun local fauna (Paleocene, Wyoming). Contributions from the Museum of Paleontology, The University of Michigan 25 (13), 259–275.

Flessa, K., Struthers, A., Fall, L.M., Dexter, T.A., 2012. Really crappy talk: fecal volume and body size in birds, mammals and dinosaurs. GSA Annual Meeting & Exposition 4–7 November 2012, Charlotte, North Carolina, USA. Paper No. 30774-206053.

Friedman, V., 2012. Vertebrate coprolites from the Lower Eagle Ford Group of north central Texas and their paleoecological significance. NMMNH Bull. 57, 221–228.


Ghosh, P., Bhattacharya, S.K., Sahni, A., Kar, R.K., Mohabey, D.M., Ambwani, K., 2003. Dinosaur coprolites from the Late Cretaceous (Maastrichtian) Lameta Formation of India: isotopic and other markers suggesting a C3 plant diet. Cretaceous Res. 24, 743–750.

Gill, F.L., Bull, I.D., 2012. Lipid analysis of vertebrate coprolites. NMMNH Bull. 57, 93–98.

Gill, F.L., Crump, M.P., Schouten, R., Bull, I.D., 2009. Lipid analysis of a ground sloth coprolite. Quaternary Res. 72, 284–288. doi: 10.1016/j.yqres.2009.06.006

Gilmore, B., 1992. Scroll coprolites from the Silurian of Ireland and the feeding of early vertebrates. Palaeontology 35, 319–333.


Habgood, K.S., Hass, H., Kerp, H., 2004. Evidence for an early terrestrial food web: coprolites from the Early Devonian Rhynie chert. T. Roy. Soc. Edin.-Earth 94, 371–389.

Hagström, J., Mehlqvist, K., 2012. The dawn of terrestrial ecosystems on Baltica: First report on land plant remains and arthropod coprolites from the Upper Silurian of Gotland, Sweden. Palaeogeogr. Palaeoclimatol. Palaeoecol. 317–318, 162–170. doi: 10.1016/j.palaeo.2012.01.001

Hansen, B.B., Milàn, J., Clemmensen, L.B., Adolfssen, J.S., Estrup, E.J., Klein, N., Mateus, O., Wings, O., 2015. Coprolites from the Late Triassic Kap Stewart Formation, Jameson Land, East Greenland: morphology, classification and prey inclusions. Geological Society, London, Special Publications, 434, 49–69. doi: 10.1144/SP434.12

Harrell, A.D., Schwimmer, D.R., 2010. Coprolites of Deinosuchus and other crocodylians from the Upper Cretaceous of Western Georgia, USA. NMMNH Bull. 51, 209–213.

Harrison, T., 2011. Coprolites: Taphonomic and Paleoecological Implications, in: Harrison, T. (Ed.), Paleontology and Geology of Laetoli: Human Evolution in Context. Volume 1: Geology, Geochronology, Paleoecology and Paleoenvironment, Vertebrate Paleobiology and Paleoanthropology. doi: 10.1007/978-90-481-9956-3_14

Hao, T.Q., Ji, C., Sun, Z.Y., Jiang, D.Y., Tintori, A., Zhang, C., 2015. Conodonts in coprolites from the Early Triassic of Chaohu, Anhui. Journal of Stratigraphy 39 (2), 65–73. [in Chinese with English abstract]

Hill, C.R., 1976. Coprolites of Ptilophyllum cuticles from the Middle Jurassic of North Yorkshire. Bull. Br. Mus. Nat. Hist. (Geol.) 27, 289–294.

Hollocher, K., Hollocher, T.C., 2012. Early processes in the fossilization of terrestrial feces to coprolites, and microstructure preservation. NMMNH Bull. 57, 79–91.

Hollocher, T.C., Chin, K., Hollocher, K.T., Kruge, M.A., 2001. Bacterial residues in coprolite of herbivorous Dinosaurs: role of bacteria in mineralization of feces. Palaios 16, 547–565.

Hollocher, K.T., Alcober, O.A., Colombi, C.E., Hollocher, T.C., 2005. Carnivore coprolites from the Upper Triassic Ischigualasto Formation, Argentina: chemistry, mineralogy, and evidence for rapid initial mineralization. Palaios 20 (1), 51–63. doi: 10.2110/palo.2003.p03-98

Hollocher, K.T., Hollocher, T.C., Rigby, J.K.Jr., 2010. A phosphatic coprolite lacking diagenetic permineralization from the Upper Cretaceous Hell Creek Formation, Northeastern Montana: importance of dietary calcium phosphate in preservation. Palaios 25 (2), 132–140. doi: 10.2110/palo.2008.p08-132r

Hone, D., Henderson, D.H., Therrien, F., Habib, M.B., 2015. A specimen of Rhamphorhynchus with soft tissue preservation, stomach contents and a putative coprolite. PeerJ 3: e1191. doi: 10.7717/peerj.1191

Hu, S.X., Zhang, Q.Y., Zhou, C.Y., 2010. Fossil coprolites from the Middle Triassic Luoping biota and ecological implication. Journal of Earth Science 21, 191–193.

Hunt, A.P., 1992. Late Pennsylvanian coprolites from the Kinney Brick Quarry, central New Mexico, with notes on the classification and utility of coprolites. Bull. New Mex. Bur. Min. Mineral Resour. 138, 221–229.

Hunt, A.P., Lucas, S.G., 2005a. A new coprolite ichnotaxon fromthe Early Permian of Texas. NMMNH Bull. 30, 121–122.

Hunt, A.P., Lucas, S.G., 2005b. A nonmarine coprolite acme zone in the Permo-Triassic. NMMNH Bull. 30, 123–124.

Hunt, A.P., Lucas, S.G., 2005c. The origin of large vertebrate coprolites from the Early Permian of Texas. NMMNH Bull. 30, 125–126.

Hunt, A.P., Lucas, S.G., 2012a. Classification of vertebrate coprolites and related trace fossils. NMMNH Bull. 57, 137–146.

Hunt, A.P., Lucas, S.G., 2012b. Descriptive terminology of coprolites and recent feces. NMMNH Bull. 57, 153–160.

Hunt, A.P., Lockley, M.G., Conrad, M.P., Paquette, M., Chure, D.J., 1993. Late Triassic vertebrates from the Dinosaur National Monument Area (Utah, USA) with an example of the utility of coprolites for correlation. NMMNH Bull. 3, 197–198.

Hunt, A.P., Chin, K., Lockley, M.G., 1994. The paleobiology of vertebrate coprolites, in: Donovan, S.K. (Ed.), The Paleobiology of Trace Fossils. John Wiley, London, pp. 221–240.

Hunt, A.P., Lucas, S.G., Lockley, M.G., 1998. Taxonomy and stratigraphic and facies significance of vertebrate coprolites of the Upper Triassic, Chinle Group, western United States. Ichnos 5, 225–234.

Hunt, A.P., Lucas, S.G., Spielmann, J.A., 2005a. Early Permian vertebrate coprolites from north-central New Mexico with description of a new ichnogenus. NMMNH Bull. 31, 39–42.

Hunt, A.P., Lucas, S.G., Spielmann, J.A., 2005b. Biochronology of Early Permian vertebrate coprolites of the American Southwest. NMMNH Bull. 31, 43–45.

Hunt, A.P., Lucas, S.G., Spielmann, J.A., Lerner, A.J., 2007. A review of vertebrate coprolites of the Triassic with descriptions of new Mesozoic ichnotaxa. NMMNH Bull. 41, 88–107.

Hunt, A.P., Lucas, S.G., Milàn, J., Spielmann, J.A., 2012a. Vertebrate coprolite studies: status and prospectus. NMMNH Bull. 57, 5–24.

Hunt, A.P., Lucas, S.G., Spielmann, J.A., 2012b. New coprolite ichnotaxa from the Buckland collection at the Oxford University Museum of Natural History. NMMNH Bull. 57, 115–124.

Hunt, A.P., Lucas, S.G., Spielmann, J.A., Cantrell, A.K., Suazo, T.L., 2012c. A new coprofauna from the Beeman Formation (Late Pennsylvanian: Late Missourian), Sacramento Mountains, New Mexico, USA. NMMNH Bull. 57, 193–196.

Hunt, A.P., Santucci, V.L., Tweet, J.S., Lucas, S.G., 2012d. Vertebrate coprolites and other bromalites in National Park Service areas. NMMNH Bull. 57, 343–353.

Hunt, A.P., Lucas, S.G., Spielmann, J.A., 2013. Triassic vertebrate coprolite ichnofaunas. NMMNH Bull. 61, 237–258.

Hunt, A.P., Lucas, S.G., Lichtig, A., 2015. A helical coprolite from the Red Crag Formation (Plio-Pleistocene) of England. NMMNH Bull. 67, 59–61.


Iñiguez, A.M., Reinhard, K.J., Araújo, A., Ferreira, L.F., Vicente, A.C.P., 2003. Enterobius vermicularis: ancient DNA from North and South American human coprolites. Mem. Inst. Oswaldo Cruz 98, 67–69.


Jain, S.L., 1983. Spirally coiled “coprolites” from the Upper Triassic Maleri Formation, India. Palaeontology 26, 813–829.


Kar, R.K., Sharma, N., Kar, R., 2004. Occurrence of fossil fungi in dinosaur dung and its implication on food habit. Curr. Sci. India 87, 1053–1056.

Kear, B.P., 2006. First gut contents in a Cretaceous sea turtle. Biology Letters 2, 113–115. doi: 10.1098/rsbl.2005.0374

Kear, B.P., Boles, W.E., Smith, E.T., 2003. Unusual gut contents in a Cretaceous ichthyosaur. Proc. R. Soc. Lond. B (Suppl.) 270, 206–208. doi: 10.1098/rsbl.2003.0050

Khosla, A., Chin, K., Alimohammadin, H., Dutta, D., 2015. Ostracods, plant tissues, and other inclusions in coprolites from the Late Cretaceous Lameta Formation at Pisdura, India: Taphonomical and palaeoecological implications. Palaeogeogr. Palaeoclimatol. Palaeoecol. 418, 90–100. doi: 10.1016/j.palaeo.2014.11.003.

Krause, J.M., Piña, C.I., 2012. Reptilian Coprolites In the Eocene of Central Patagonia, Argentina. J. Paleontol. 86 (3), 527–538.


Laojumpon, C., Matkhammee, T., Wathanapitaksakul, A., Suteethorn, V., Suteethorn, S., Lauprasert, K., Srisuk, P., Loeuff, J.L., 2012. Preliminary report on coprolites from the Late Triassic of Thailand. NMMNH Bull. 57, 207–213.

Linseele, V., Riemer, H., Baeten, J., De Vos, D., Marinova, E., Ottoni, C., 2013. Species identification of archaeological dung remains: A critical review of potential methods. Journal of Environmental Archaeology 18 (1), 5–17.  doi: 10.1179/1461410313Z.00000000019

Lockley, M.G., Gierlinski, G.D., Houck, K., Lim, J.D., Kim, K.S., Kim, D.Y., Kim, T.Y., Kang, S.H., Foster, R.H., Li, R., Chessser, C., Gay, R., Dubicka, Z., Cart, K., Wright, C., 2014. New excavation at the Mill Canyon dinosaur track site (Cedar Mountain Formation, Lower Cretaceous) of eastern Utah. NMMNH Bull. 62, 287–300.

Lomax, D.R., 2010. An Ichthyosaurus (Reptilia, Ichthyosauria) with gastric contents from Charmouth, England: first report of the genus from the Pliensbachian. Paludicola 8 (1), 22–36.

Long, C., Wings, O., Xiaohong, C., Sander, P.M., 2006. Gastroliths in the Triassic ichthyosaur Panjiangsaurus from China. J. Paleont. 80 (3), 583–588.

Lucas, S.G., Spielmann, J.A., Hunt, A.P., Emry, R.J., 2012. Crocodylian coprolites from the Eocene of the Zaysan basin, Kazakstan. NMMNH Bull. 57, 319–324.

Luo, M., Hud, S., Benton, M.J., Shi, G.R., Zhao, L., Huang, J., Song, H., Wen, W., Zhang, Q., Fang, Y., Huang, Y., Chen, Z.Q., 2016. Taphonomy and palaeobiology of early Middle Triassic coprolites from the Luoping biota, southwest China: Implications for reconstruction of fossil food webs. Palaeogeogr. Palaeoclimatol. Palaeoecol. 474, 232–246. doi: 10.1016/j.palaeo.2016.06.001


Mancuso, A.C., Marsicano, C., Palma, R., 2004. Vertebrate coprolites from the Triassic of Argentina (Cuyana Basin). Ameghiniana 41 (3), 347–354.

Martínez, A.C., Yagueddú, C., 2012. Identificación de microrrestos vegetales en un coprolito humano del sitio Cerro Casa de Piedra, Santa Cruz, Argentina. Magallania 40 (1), 333–339.

McAllister, J.A., 1985. Reevaluation of the formation of spiral coprolites. The University of Kansas Paleontological Contributions 114, 1–12.

Milàn, J., Clemmensen, L.B., Schulz Adolfssen, J., Estrup, E.J., Frobøse, N., Klein, N., Mateus, O., Wings, O., 2012. A preliminary report on coprolites from the Late Triassic part of the Kap Stewart Formation, Jameson Land, east Greenland. NMMNH Bull. 57, 203–205.

Milàn, J., Rasmussen, B.W., Lynnerup, N., 2012. A coprolite in the MDCT-scanner – internal architecture and bone contents revealed. NMMNH Bull. 57, 99–103.

Milàn, J., Rasmussen, B., Bonde, N., 2012. Coprolites with prey remains and traces from coprophagous organisms from the Lower Cretaceous (Late Berriasian) Jydegaard Formation of Bornholm, Denmark. NMMNH Bull. 57, 235–240.

Milàn, J., Hunt, A.P., Adolfssen, J.S., Rasmussen, B.W., Morten, B., 2015. First record of a vertebrate coprolite from the Upper Cretaceous (Maastrichtian) chalk of Stevns Klint, Denmark. NMMNH Bull. 67, 227–229.

Monaco, P., Baldanza, A., Bizzarri, R., Famiani, F., Lezzerini, M., Sciuto, F., 2014. Ambergris cololites of Pleistocene sperm whales from central Italy and description of the new ichnogenus and ichnospecies Ambergrisichnus alleronae. Palaeontologia Electronica 17 (2), 1–20.

Mustoe, G.E., 2001. Enigmatic origin of ferruginous „coprolites”: Evidence from the Miocene wilkes Formation, southwestern Washington. Geol. Soc. Am. Bull. 113, 673–681.


Nakajima, Y., Izumi, K., 2014. Coprolites from the upper Osawa Formation (upper Spathian), northeastern Japan: Evidence for predation in a marine ecosystem 5 Myr after the end-Permian mass extinction. Palaeogeogr. Palaeoclimatol. Palaeoecol. 414, 225–232. doi: 10.1016/j.palaeo.2014.08.014.

Neto de Carvalho, C., Farinha, C., 2006. Coprolites From Portugal: A Synthesis With The Report Of New Findings. Ichnology Newsletter 27, 10–15.

Niedźwiedzki, G., 2011. Dinosaur tracks from the Early Jurassic ecosystem of Sołtyków, Holy Cross Mountains. Biuletyn Państwowego Instytutu Geologicznego 447, 49–98. [in Polish with English abstract and summary]

Niedźwiedzki, G., Bajdek, P., Qvarnström, M., Sulej, T., Sennikov, A.G., Golubev, V.K., 2016. Reduction of vertebrate coprolite diversity associated with the end-Permian extinction event in Vyazniki region, European Russia. Palaeogeogr. Palaeoclimatol. Palaeoecol. 450, 77–90. doi: 10.1016/j.palaeo.2016.02.057

Niedźwiedzki, G., Bajdek, P., Owocki, K., Kear, B.P., 2016. An Early Triassic polar predator ecosystem revealed by vertebrate coprolites from the Bulgo Sandstone (Sydney Basin) of southeastern Australia. Palaeogeogr. Palaeoclimatol. Palaeoecol. 464, 5–15. doi: 10.1016/j.palaeo.2016.04.003

Northwood, C., 2005. Early Triassic coprolites from Australia and their palaeobiological significance. Palaeontology 48, 49–68.


O’Keefe, F.R., Street, H.P., Cavigelli, J.P., Socha, J.J., O’Keefe, R.D., 2009. A plesiosaur containing an ichthyosaur embryo as stomach contents from the Sundance Formation of the Bighorn basin, Wyoming. J. Vertebr. Paleontol. 29 (4), 1306–1310.

Owocki, K., Niedźwiedzki, G., Sennikov, A.G., Golubev, V.K., Janiszewska, K., Sulej, T., 2012. Upper Permian vertebrate coprolites from Vyazniki and Gorokhovets, Vyatkian regional stage, Russian Platform. Palaios 27, 867–877. doi: palo.2012.p12-017r


Poinar, G., Boucot, A.J., 2006. Evidence of intestinal parasites of dinosaurs. Parasitology 133, 245–249.

Prasad, V., Strömberg, C.A.E., Alimohammadian, H., Sahni, A., 2005. Dinosaur coprolites and the early evolution of grasses and grazers. Science 310, 1177–1180.

Přikryl, T., Košťák, M., Mazuch, M., Mikuláš, R., 2012. Evidence for fish predation on a coleoid cephalopod from the Lower Jurassic Posidonia Shale of Germany. N. Jb. Geol. Paläont. Abh. 263 (1), 25–33.


Qvarnström, M., Niedźwiedzki, G., Žigaitė, Ž., 2016. Vertebrate coprolites (fossil faeces): An underexplored Konservat-Lagerstätte. Earth-Science Reviews 162, 44–57. doi: 10.1016/j.earscirev.2016.08.014


Retallack, G.J., Krull, E.S., 1997. Permian coprolites from Graphite Peak, Antarctica. U.S. Antarctic Journal. 30 (5), 7–9.

Robbins, E.I., Porter, K.G., Haberyan, K.A., 1985. Pellet microfossils: Possible evidence for metazoan life in Early Proterozoic time. Proc. Natl. Acad. Sci. USA 82, 5809–5813.

Rodríguez-de la Rosa, R.A., Cevallos-Ferriz, S.R.S., Silva-Pineda, A., 1998. Paleobiological implications of Campanian coprolites. Palaeogeogr. Palaeoclimatol. Palaeoecol. 142, 231–254.

Rothwell, G.W., Scott, A.C., 1983. Coprolites within marattiaceous fern stems (Psaronius magnificus) from the upper Pennsylvanian of the Appalachian Basin, U.S.A. Palaeogeogr. Palaeoclimatol. Palaeoecol. 41, 227–232. doi: 10.1016/0031-0182(83)90089-5

Ruiz-Omeñaca, J.I., Canudo, J.I., Cuenca-Bescós, G., 1997. First evidence of an herbivorous dinosaurs feeding area in the Lower Cretaceous of Spain (Teruel). Monografías de la Academia de Ciencias Exactas, Físicas, Químicas y Naturales de Zaragoza 10, 1–48. [in Spanish with English abstract]


Santoro, C., Vinton, S.D., Reinhard, K.J., 2003. Inca expansion and parasitism in the Lluta Valley: preliminary data. Mem. Inst. Oswaldo Cruz 98, 161–163.

Schweigert, G., Dietl, G., 2012. Vertebrate coprolites from the Nusplingen Lithographic Limestone (Upper Jurassic, SW Germany). NMMNH Bull. 57, 215–220.

Schwimmer, D.R., Weems, R.E., Sanders, A.E., 2015. A Late Cretaceous shark coprolite with baby freshwater turtle vertebrae inclusions. Palaios 30, 707–713. doi: 10.2110/palo.2015.019

Scott, A.C., 1977. Coprolites containing plant material from the Carboniferous of Britain. Palaeontology 20 (1), 59–68.

Scott, A.C., Taylor, T.N., 1983. Plant/animal interactions during the Upper Carboniferous. Bot. Rev. 49 (3), 259–307. doi: 10.1007/BF02861089

Scott, A.C., Chaloner, W.G., Paterson, S., 1985. Evidence of pteridophyte–arthropod interactions in the fossil record. P. Roy. Soc. Edinb. B 86, 133–140. doi: 10.1017/S0269727000008058

Sharma, N., Kar, R.K., Agarwal, A., Kar, R., 2005. Fungi in dinosaurian (Isisaurus) coprolites from the Lameta Formation (Maastrichtian) and its reflection on food habit and environment. Micropaleontology 51 (1), 73–82.

Segesdi, M., Botfalvai, G., Bodor, E.R., Ösi, A., Buczkó, K., Dallos, Z., Tokai, R., Földes, T., 2017. First report on vertebrate coprolites from the Upper Cretaceous (Santonian) Csehbánya Formation of Iharkút, Hungary. Cretaceous Research 74, 87–99. doi: 10.1016/j.cretres.2017.02.010

Seilacher, A., Marshall, C., Skinner, H.C.W., Tsuihiji, T., 2001. A fresh look at sideritic “coprolites”. Paleobiology 27 (1), 7–13.

Shelton, C.D., 2013. A new method to determine volume of bromalites: morphometrics of Lower Permian (Archer City Formation) heteropolar bromalites. Swiss J. Palaeontol. 132, 221–238. doi: 10.1007/s13358-013-0057-z

Silva, R.R., Ferigolo, J., Bajdek, P., Piñeiro, G., 2017. The feeding habits of Mesosauridae. Front. Earth Sci. 5:23. doi: 10.3389/feart.2017.00023

Sistiaga, A., Berna, F., Laursen, R., Goldberg, P., 2014. Steroidal biomarker analysis of a 14,000 years old putative human coprolite from Paisley Cave, Oregon. Journal of Archaeological Science 41, 813–817. doi: 10.1016/j.jas.2013.10.016

Sistiaga A., Mallol, C., Galván, B., Summons, R.E., 2014. The Neanderthal Meal: A new perspective using faecal biomarkers. PLOS ONE 9 (6), e101045. doi: 10.1371/journal.pone.0101045

Smith, R.M.H., Botha-Brink, J., 2011. Morphology and composition of bone-bearing coprolites from the Late Permian Beaufort Group, Karoo Basin, South Africa. Palaeogeogr. Palaeoclimatol. Palaeoecol. 312, 40–53. doi: 10.1016/j.palaeo.2011.09.006

Souto, P.R.F., 2001. Tetrapod coprolites from the Middle Triassic of southern Brazil. Gaia 16, 51–57.

Souto, P.R.F., 2010. Crocodylomorph coprolites from the Bauru basin, Upper Cretaceous, Brazil. NMMNH Bull. 51, 201–208.

Stewart, J.D., 1978. Enterospirae (fossil intestines) from the Upper Cretaceous Niobrara Formation of Western Kansas. The University of Kansas Paleontological Contributions 89, 9–16.

Stone, D.D., Crisp, E.L., Bishop, J.T., 2000. A large meat-eating dinosaur coprolite from the Jurassic Morrison Formation of Utah. Geol. Soc. Am. Abstr. Programs 32 (7), 220.

Suazo, T.L., Cantrell, A.K., Lucas, S.G., Spielmann, J.A., Hunt, A.P., 2012. Coprolites across the Cretaceous/Tertiary boundary, San Juan Basin, New Mexico. NMMNH Bull. 57, 263–274.

Sullivan, R.M., Jasinski, S.E., 2012. Coprolites from the Upper Cretaceous Fruitland, Kirtland and Ojo Alamo formations, San Juan basin, New Mexico. NMMNH Bull. 57, 255–262.


Teixeira-Santos, I., Sianto, L., Araújo, A., Reinhard, K.J., Chaves, S.A.M., 2015. The evidence of medicinal plants in human sediments from Furna do Estrago prehistoric site, Pernambuco State, Brazil. Quatern. Int. 377, 112–117. doi: 10.1016/j.quaint.2015.01.019

Thies, D., Hauff, R.B., 2013. A Speiballen from the Lower Jurassic Posidonia Shale of South Germany. N. Jb. Geol. Paläont. Abh. 267 (1), 117–124.

Thulborn, R.A., 1991. Morphology, preservation and palaeobiological significance of dinosaur coprolites. Palaeogeogr. Palaeoclimatol. Palaeoecol. 83, 341–366.

Tweet, J.S., Chin, K., Braman, D.R., Murphy, N.L., 2008. Probable gut contents within a specimen of Brachylophosaurus canadensis (Dinosauria: Hadrosauridae) from the Upper Cretaceous Judith River Formation of Montana. Palaios 23, 624–635. doi: 10.2110/palo.2007.p07-044r

Tweet, J., Chin, K., Ekdale, A.A., 2016. Trace fossils of possible parasites inside the gut contents of a hadrosaurid dinosaur, Upper Cretaceous Judith River Formation, Montana. J. Paleontol. 90 (2), 279–287. doi: 10.1017/jpa.2016.43


van Geel, B., Guthrie, R.D., Altmann, J.G., Broekens, P., Bull, I.D., Gill, F.L., Jansen, B., Nieman, A.M., Gravendeel, B., 2011. Mycological evidence of coprophagy from the feces of an Alaskan Late Glacial mammoth. Quaternary Sci. Rev. 30 (17), 2289–2303. doi: 10.1016/j.quascirev.2010.03.008

van Geel, B., Protopopov, A., Bull, I., Duijm, E., Gill, F., Lammers, Y., Nieman, A., Rudaya, N., Trofimova, S., Tikhonov, A.N., Vos, R., Zhilich, S., Gravendeel, B., 2014. Multiproxy diet analysis of the last meal of an early Holocene Yakutian bison. J. Quaternary Sci. 29 (3), 261–268. doi: 10.1002/jqs.2698

Vannier, J., Chen, J., 2005. Early Cambrian Food Chain: New Evidence from Fossil Aggregates in the Maotianshan Shale Biota, SW China. Palaios 20, 3–26. doi: 10.2110/palo.2003.p03-40

Verde, M., and Ubilla, M. 2002. Mammalian carnivore coprolites from the Sopas Formation (Upper Pleistocene, Lujanian Stage), Uruguay. Ichnos 9 (1): 77–80. doi: 10.1080/10420940190034102

Vijaya, G., Prasad, V.R., Singh, K., 2009. Late Triassic palynoflora from the Pranhita–Godavari Valley, India: evidence from vertebrate coprolites. Alcheringa 33 (2), 91–111. doi: 10.1080/03115510902841315

Vinton, S.D., Perry, L., Reinhard, K.J., Santoro, C.M., Teixeira-Santos, I., 2009. Impact of empire expansion on household diet: the Inka in Northern Chile’s Atacama Desert. PloS ONE, 4 (11), e8069. doi: 10.1371/journal.pone.0008069


Wahl, A.M., Martin, A.J., Hasiotis, S., 1998. Vertebrate coprolites and coprophagy traces, Chinle Formation (Late Triassic), Petrified Forest National Park, Arizona, in: Santucci, V.L., McClelland, L. (Eds.), National Park Service Paleontological Research. Lakewood, CO: National Park Service Geological Resources Division Technical Report NPS/NRGRD/GRDTR-98/01, pp. 144–148.

Wang, M., Zhou, Z., Sullivan, C., 2016. A Fish-Eating Enantiornithine Bird from the Early Cretaceous of China Provides Evidence of Modern Avian Digestive Features. Curr. Biol. 26, 1–7. doi: 10.1016/j.cub.2016.02.055

Williams, M.E., 1972. The origin of “spiral coprolites”. The University of Kansas Paleontological Contributions 59, 1–19.

Wings, O., 2007. A review of gastrolith function with implications for fossil vertebrates and a revised classification. Acta Palaeontol. Pol. 52 (1), 1–16.

Wings, O., 2012. Gastroliths in coprolites — a call to search! NMMNH Bull. 57, 73–77.

Wings, O., 2015. The rarity of gastroliths in sauropod dinosaurs – a case study in the Late Jurassic Morrison Formation, western USA. Foss. Rec. 18, 1–16. doi: 10.5194/fr-18-1-2015

Wood, J.R., 2007. Moa gizzard content analyses: further information on the diets of Dinornis robustus and Emeus crassus, and the first evidence for the diet of Pachyornis elephantopus (Aves: Dinornithiformes). Records of the Canterbury Museum 21, 27–39.

Wood, J.R., Wilmshurst, J.M., 2013. Pollen analysis of coprolites reveals dietary details of heavy-footed moa (Pachyornis elephantopus) and coastal moa (Euryapteryx curtus) from Central Otago. New Zealand Journal of Ecology 37 (1), 151–155.

Wood, J.R., Wilmshurst, J.M., 2014. Late Quaternary terrestrial vertebrate coprolites from New Zealand. Quaternary Sci. Rev. 98, 33–44. doi: 10.1016/j.quascirev.2014.05.020

Wood, J.R., Wilmshurst, J.M., 2016. A protocol for subsampling Late Quaternary coprolites for multi-proxy analysis. Quaternary Sci. Rev. 138, 1–5. doi: 10.1016/j.quascirev.2016.02.018

Wood, J.R., Rawlence, N.J., Rogers, G.M., Austin, J.J., Worthy, T.H., Cooper, A., 2008. Coprolite deposits reveal the diet and ecology of the extinct New Zealand megaherbivore moa (Aves, Dinornithiformes). Quaternary Science Reviews 27, 2593–2602. doi: 10.1016/j.quascirev.2008.09.019

Wood, J.R., Wilmshurst, J.M., Wagstaff, S.J., Worthy, T.H., Rawlence, N.J., Cooper, A., 2012. High-Resolution Coproecology: Using Coprolites to Reconstruct the Habits and Habitats of New Zealand’s Extinct Upland Moa (Megalapteryx didinus). PLoS ONE 7, e40025. doi: 10.1371/journal.pone.0040025

Wood, J.R., Wilmshurst, J.M., Worthy, T.H., Cooper, A., 2012. First coprolite evidence for the diet of Anomalopteryx didiformis, an extinct forest ratite from New Zealand. New Zealand Journal of Ecology 36 (2), 164–170.

Wood, J.R., Wilmshurst, J.M., Rawlence, N.J., Bonner, K.I., Worthy, T.H., Kinsella, J.M., Cooper, A., 2013. A Megafauna’s microfauna: gastrointestinal parasites of New Zealand’s extinct moa (Aves: Dinornithiformes). PLoS ONE 8 (2), e57315. doi: 10.1371/journal.pone.0057315

Wood, J.R., Wilmshurst, J.M., Richardson, S.J., Rawlence, N.J., Wagstaff, S.J., Worthy, T.H., Cooper, A., 2013. Resolving lost herbivore community structure using coprolites of four sympatric moa species (Aves: Dinornithiformes). PNAS 110 (42), 16910–16915.


Yates, A.M., Neumann, F.H., Hancox, P.J., 2012. The Earliest Post-Paleozoic Freshwater Bivalves Preserved in Coprolites from the Karoo Basin, South Africa. PLoS ONE 7 (2), e30228. doi: 10.1371/journal.pone.0030228


Zatoń, M., Rakociński, M., 2014. Coprolite evidence for carnivorous predation in a Late Devonian pelagic environment of southern Laurussia. Palaeogeogr. Palaeoclimatol. Palaeoecol. 394, 1–11. doi: 10.1016/j.palaeo.2013.11.019

Zatoń, M., Niedźwiedzki, G., Marynowski, L., Benzerara, K., Pott, C., Cosmidis, J., Krzykawski, T., Filipiak, P., 2015. Coprolites of Late Triassic carnivorous vertebrates from Poland: An integrative approach. Palaeogeogr. Palaeoclimatol. Palaeoecol. 430, 21–46. doi: 10.1016/j.palaeo.2015.04.009

Zatoń M., Broda K., Qvarnström M., Niedźwiedzki G., Ahlberg P.E., 2017. The first direct evidence on a Late Devonian coelacanth fish feeding on conodont animals. Sci. Nat. 104:26. doi: 10.1007/s00114-017-1455-7

Zhou, Z., Clarke, J., Zhang, F., Wings, O., 2004. Gastroliths in Yanornis: an indication of the earliest radical diet-switching and gizzard plasticity in the lineage leading to living birds? Naturwissenschaften 91, 571–574. doi: 10.1007/s00114-004-0567-z

Zítt, J., Vodrázka, R., Hradecká, L., Svobodová, M., Stastný, M., Svábenická, L., 2015. Depositional and palaeoenvironmental variation of lower Turonian nearshore facies in the Bohemian Cretaceous Basin, Czech Republic. Cretaceous Res. 56, 293–315. doi: 10.1016/j.cretres.2015.05.007

Modern feces


Balazs, G.H., Fujioka, R., Fujioka, C., 1993. Marine Turtle Faeces on Hawaiian Beaches. Mar. Pollut. Bull. 26 (7), 392–394. doi: 10.1016/0025-326X(93)90187-O

Barnes, R.F.W., 2001. How reliable are dung counts for estimating elephant numbers? Afr. J. Ecol. 39, 1–9.

Beyer, W.N., Connor, E.E., Gerould, S., 1994. Estimates of soil ingestion by wildlife. J. Wildl. Manag. 58 (2), 375–382.


Chame, M., 2003. Terrestrial mammal feces: a morphometric summary and description. Mem. Inst. Oswaldo Cruz 98 (Suppl. I), 71–94.

Chame, M., Sianto, L., 2011. Diagnóstico de Coprólitos, ou Quem Fez o Coprólito? In: Ferreira, L.F., Reinhard, K., Araújo, A. (Eds.), Fundamentos da Paleoparasitologia. Editora Fiocruz, pp. 263–281. [in Portuguese]


Dauphin, Y., Andrews, P., Denys, C., Fernández-Jalvo, Y., Williams, T., 2003. Structural and chemical bone modifications in a modern owl pellet assemblage from Olduvai Gorge (Tanzania). Journal of Taphonomy 1 (4), 209–232.

Davison, A., Birks, J.D.S., Brookes, R.C., Braithwaite, T.C., Messenger, J.E., 2002. On the origin of faeces: morphological versus molecular methods for surveying rare carnivores from their scats. J. Zool. (Lond.) 257, 141–143.


Esque, T.C., Peters, E.L., 1994. Ingestion of bones, stones, and soil by desert tortoises. Fish and Wildlife Researchers 13, 105–111.


Galante, E., Marcos-García, M.A., 1997. Detritívoros, coprófagos y necrófagos. Los Artrópodos y el Hombre 20, 57–64. [in Spanish]

Gill, F.L., Dewhurst, R.J., Evershed, R.P., McGeough, E., O’Kiely, P., Pancost, R.D., Bull, I.D., 2011. Analysis of archaeal ether lipids in bovine faeces. Anim. Feed Sci. Tech. 87–92. doi:10.1016/j.anifeedsci.2011.04.006

Gupta, M., Aggarwal, P., Singh, R., Lehl, S.S, 2014. A Case of Giant Fecaloma in a 32-Year-Old Woman. Austin J. Clin. Case Rep. 1 (4), id1017.


Halfpenny, J.C., 2008. Scats and tracks of North America: a field guide to the signs of nearly 150 wildlife species. Falcon Guides, USA, 336 pp.


Keenan, S.W., Engel, A.S., Elsey, R.M., 2013. The alligator gut microbiome and implications for archosaur symbioses. Scientific Reports 3, 2877. doi: 10.1038/srep02877


Lewis, S.J., Heaton, K.W., 1997. Stool form as a useful guide to intestinal transit time. Scand. J. Gastroenterol. 32, 920–924.

Lock, B., Heard, D., Detrisac, C., Elliott Jacobson, E., 2003. An epizootic of chronic regurgitation associated with chlamydophilosis in recently imported emerald tree boas (Corallus caninus). J. Zoo. Wildlife Med. 34 (4), 385–393.


Milàn, J., 2012. Crocodylian scatology – a look into morphology, internal architecture, inter- and intraspecific variation and prey remains in extant crocodylian feces. NMMNH Bull. 57, 65–71

Mumma, M.A., Adams, J.R., Zieminski, C., Fuller, T.K., Mahoney, S.P., Waits, L.P., 2015. A comparison of morphological and molecular diet analyses of predator scats. J. Mammal. doi: 10.1093/jmammal/gyv160


Norris, D.O., Bock, J.H., 200. Use of fecal material to associate a suspect with a crime scene: report of two cases. J. Forensic Sci. 45 (1), 184–187.


Parameshwarappa, K.D., Chandrakanth, C., Sunil, B., 2012. The prevalence of intestinal parasitic infestations and the evaluation of different concentration techniques of the stool examination. Journal of Clinical and Diagnostic Research 6 (7), 1188-1191.

Pauwels, O.S.G., Barr, B., Sanchez, M.L., Burger, M., 2007. Diet records for the dwarf crocodile, Osteolaemus tetraspis tetraspis in Rabi Oil Fields and Loango National Park, southwestern Gabon. Hamadryad 31 (2), 258–264.


Sistiaga, A., Wrangham, R., Rothman, J.M., Summons, R.E., 2015. New insights into the evolution of the human diet from faecal biomarker analysis in wild chimpanzee and gorilla faeces. PLoS ONE 10 (6), e0128931. doi: 10.1371/journal.pone.0128931


Wasser, S.K., Houston, C.S., Koehler, G.M., Cadd, G.G., Fain, S.R., 1997. Techniques for application of faecal DNA methods to field studies of Ursids. Mol. Ecol. 6, 1091–1097.


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