Paleoecological and taphonomical aspects of the Late Miocene vertebrate coprolites (Urumaco Formation) of Venezuela

Abstract

Coprolites (fossil feces) provide important paleoecological information, such as diet of the producer, parasite infestation and gut microbiota, which cannot be accessed through body remains. The Upper Miocene Urumaco Formation, in western Venezuela, has a diverse vertebrate fauna composed of ground sloths, rodents, reptiles, fishes and amphibians, as well as coprolites. Here we describe 106 coprolites produced by a range of carnivorous and herbivorous vertebrates, classified into five different morphotypes based on their size and shape, suggesting different producers. Herbivorous morphotypes (M1, M2 and M5) are the most abundant (59%). The most common inclusions in these coprolites are Poaceae remains, although Eudicotyledoneae fragments, bacteria and cyanobacteria filaments were also found. The M1 morphotype has a bullet shape with longitudinal grooves; the M2 morphotype is rounded to oval, with at least one distinctive tapered end; and the M5 morphotype is a short cylinder with a rough surface, and rounded ends. In contrast, the morphotypes M3 and M4 are attributed by us to carnivores and have different inclusions. While M3 is a large cylinder with rounded ends and contains bacteria, nematode eggs and phytoliths, M4 is sausage-shaped and has muscle cells preserved inside. Based on the co-occurring fauna and the morphology of the coprolites, the probable producers were inferred as the following: M1, a rodent; M2, a notoungulate; M3, a crocodilian; M4, an indeterminate carnivore; and M5, a sirenian. The preservation of herbivorous and carnivorous coprolites with bacterial, parasitological and muscle remains in the same formation and even the same outcrop is rare in the geological record. This unique association and preservation suggests early lithification promoted by rapid burial, although the presence of the boring trace Gastrochaenolites in a single specimen indicates some degree of time-averaging.