Vertebrate coprolites

Buckland introduced the term coprolite for fossilized feces in 1829, and there is an extensive literature on coprolites. Latrinites (accretionary and ethological) are accumulations of coprolites. Clusters of conodont elements from the Early Ordovician probably represent coprolites produced by conodont animals, which, for those who consider conodonts to be vertebrates, are the oldest vertebrate coprolites. Vertebrate coprolites are common in the Devonian, with the most important assemblages derived from fossil-fish localities in Scotland. Carboniferous strata record the first large samples of vertebrate coprolites, and the first abundant samples occur in the Pennsylvanian. The Carboniferous coprolite record is strictly Laurussian, and the Permian coprolite record globalizes to include records from Gondwana. The Late Triassic sample of coprolites is by far the largest from this period, and the majority of specimens are from nonmarine strata. Dung fungus Sporormiella provides a proxy for the introduction of grazing herbivores. The majority of known human coprolite sites are from North America. The description and naming of vertebrate coprolites is important and has practical applications in detailed analyses of distribution patterns, including a robust biostratigraphy and a biochronology for the Triassic, utilizing 14 ichnogenera. Ichnotaxobases for coprolites include (1) external morphology; (2) internal structure; (3) surface texture; (4) size; (5) inclusions; and (6) mineralogy. We recognize 56 valid ichnotaxa of vertebrate coprolites. Producer of coprolites can be identified by morphology, analogy, association in digest tract, association with skeletal remains, biochemistry, inclusions, and size. Coprolites can serve as proxies for biotaxa (the animals that produced them). The producer of individual vertebrate coprolites can usually only be identified at a high taxonomic level (order or higher) with very few exceptions before the Pleistocene. Vertebrate coprolites provide some biochronology in the Pennsylvanian to the Quaternary. Vertebrate coprolites can delimit biogeographic regions. All trace fossils, including coprolites, are facies fossils. Spiral vertebrate coprolites in shallow marine environments define the Crassocoprus ichnofacies that ranges from the Mississippian to the Eocene. On a finer scale, vertebrate coprolites allow the discrimination of three ichnocoenoses in the nonmarine Late Triassic and four in an ecological transect from lacustrine to shallow marine conditions in the Late Pennsylvanian. Some coprolites represent Lagerstätten. Coprolites can provide diverse information about the diet of their producers and contribute to evidence of digestive processes. Coprolites have long been recognized as having utility in the reconstructions of ancient trophic relations. The analysis of human coprolites can address health issues. Coprolites can also be used as biotaxa proxies to analyze ecosystem evolution, extinction, and diversity patterns.