Trace fossil evidence for infaunal moulting in a Middle Devonian non-trilobite euarthropod

Trace fossils represent the primary source of information on the evolution of animal behaviour through deep time, and provide exceptional insights into complex life strategies that would be otherwise impossible to infer from the study of body parts alone. Here, we describe unusual trace fossils found in marginal-marine, storm- and river-flood deposits from the Middle Devonian Naranco Formation of Asturias (northern Spain) that constitute the first evidence for infaunal moulting in a non-trilobite euarthropod. The trace fossils are preserved in convex hyporelief, and include two main morphological variants that reflect a behavioural continuum. Morphotype 1 consists of a structure that superficially resembles a Rusophycus with an oval outline that possesses a distinctly three lobed axis with an elevated central ridge and regularly spaced transverse furrows that convey the appearance of discrete body segments. The anterior part is the most irregular region of the structure, and it is not always recorded. Morphotype 2 displays more elongated, tubular morphology. Careful observation, however, reveals that it comprises up to three successive morphotype 1 specimens organised in a linear fashion and partially truncating each other. Trilobate morphology and effaced transverse furrows are locally evident, but the predominant morphological feature is the continuous, elevated ridge. The detailed morphology of morphotype 1 and well-preserved, discrete segments of morphotype 2 closely resemble the dorsal exoskeleton of the enigmatic late Carboniferous euarthropod Camptophyllia, suggesting the possible affinities of the producer. Comparisons with patterns of Devonian phacopid trilobite exuviation suggest that the Naranco Formation trace fossils may have been produced by the infaunal activities of an euarthropod that anchored its dorsal exoskeleton in the firm sediment during the body inversion moult procedure. Our findings expand the phylogenetic and environmental occurrence of infaunal moulting in Palaeozoic euarthropods, and suggest a defensive strategy against predation, previously only known from trilobites preserved in open-marine deposits.