First evidence of reproductive strategies in cephalopods preserved in phosphate and siderite nodules from the Devonian of Uruguay

Abstract

Uruguayan ammonoids are preserved in phosphate and siderite nodules found at the basalmost tillite-like conglomerates of the San Gregorio Formation. This lithostratigraphic unit was deposited under glacial conditions and its age (as well as that of the nodules) has been highly debated because glaciations were intermittent in Gondwana during the Late Paleozoic. Reef-builder organisms (e.g., Rugosa and Tabulata), goniatite and orthoceratid cephalopods, brachiopods, sponges, actinopterygians and other indeterminate gnatostomes, as well as fragmentary stems and roots of cf. Lycopsida are the most frequent fossils in the nodules. According to new biostratigraphic and paleoclimatic evidence, these taxa are representative of a reefal environment of a preliminary Devonian age including species that are common in the underlying Early Devonian (Emsian) Durazno Group. Among the ammonoid remains, more than 40 clusters of hatchling goniatites were found in the nodules. Each cluster contains a variable number of shells similar in shape to some of the adults also preserved within the nodules, representing a single species preserved at the same developmental stage (3 mm on size average). The strongly packed shells are enveloped by a substance with a different chemical composition and microstructure with respect to that of the nodule matrix, possibly indicating the presence of a gelatinous-like substance reminiscent of that secreted by the females of some extant cuttlefish and octopuses at the time of the egg spawn. Differing from previously described ammonoid accumulations, our clusters are unique in containing individuals of just a single species preserved in the same ontogenetic stage. That allows us to suggest that they represent a mode of reproduction in which hatchlings were morphologically similar to their parents and occupied the same habitat. Our results are thus one of the oldest known records of reproductive strategies in Paleozoic ammonoids and the phosphate and siderite nodules from the San Gregorio Formation are here classified as a new Konservat-Lagerstätte, which is the oldest known for South America.