Evidence for an early terrestrial food web: coprolites from the Early Devonian Rhynie chert

Abstract

Fossils in the Pragian Rhynie cherts, Aberdeenshire, Scotland, are preserved with exquisite cellular detail, and provide much information on Early Devonian terrestrial and freshwater ecosystems. The fossils include abundant and diverse coprolites which demonstrate the existence of consumers differing in life-habit and diet. The coprolites are small (0.5–3 mm) and diverse in morphology and content, including groups of amorphous coprolites as well as coprolites with identifiable, particulate content. The present authors define three new ichnogenera to accommodate these coprolites:

• • Lancifaex encompasses elongate coprolites with particulate content, and includes three ichnospecies, distinguished on morphology, L. simplex, L. divisa and L. moniliforma.

• • Rotundafaex encompasses rotund coprolites with particulate content, and includes a single ichnospecies, R. aggregata.

• • Bacillafaex encompasses rod-shaped coprolites with amorphous content, and includes two ichnospecies, distinguished on size, B. constipatus and B. mina.

Typically, plant spores do not dominate the content of these coprolites, but the population does include some spore-rich coprolites. The presence of spore-rich coprolites in this diverse assemblage adds evidence to the debate on spore-feeding as a nutritional strategy in early terrestrial ecosystems. The authors conclude that the coprolites described here indicate at least four types of consumer including detritivores and herbivores. Spore-rich coprolites might suggest sporivory; however, comparison with the faeces of modern millipedes demonstrates that they might equally well be the product of detritivores. The continuum observed here between spore-rich and spore-poor coprolites implies that, in this assemblage, spore-rich coprolites do not constitute a distinct group. Rather, they are part of a group of elongate, ellipsoidal coprolites with heterogeneous content that includes, at one extreme, coprolites which lack plant spores, and at the other, coprolites which contain abundant plant spores. Most coprolites in this group fall somewhere between the two extr