Heliolitid corals and their competitors: a case study from the Wellin patch reefs, Middle Devonian, Belgium

peer reviewedWellin patch reefs are small Upper Eifelian build?ups within the fine?grained argillaceous limestone of the Hanonet Formation. Whereas the reefs themselves are not well exposed, their fossil assemblage is accessible in the hills near the town of Wellin, approximately 40xA0km SE of Dinant in Belgium. It is especially rich in massive stromatoporoids, heliolitids and other tabulate corals. They exhibit predominantly domical and bulbous morphologies. This paper focuses primarily on the palaeoautoecology of the heliolitid corals and their relationships with other organisms. Cases of mutual overgrowth between heliolitids, other corals and stromatoporids suggest a high degree of competition for space on the reefs, possibly related to the scarcity of hard substrates. Coral and stromatoporoid growth forms, as well as the prevalence of micritic matrix, point to a relatively low energy environment. However, abundant growth interruption surfaces, sediment intercalations and rejuvenations of corals suggest episodically increased hydrodynamic regime and sediment supply. It is inferred that the patch reefs developed in a relatively shallow environment, where the reefal assemblage was regularly affected by storms. Heliolitids exhibited high sediment tolerance and relied on passive sediment removal for survival. They also could regenerate effectively and commonly overgrew their epibionts, after the colony’s growth was hampered by the sediment. This is recorded in extremely abundant growth interruption surfaces, which allow the analysis of the impact of sediment influxes on the heliolitid corals. ? 2021 Lethaia Foundation. Published by John Wiley & Sons Lt

Heliolitid corals and their competitors: a case study from the Wellin patch reefs, Middle Devonian, Belgium

peer reviewedWellin patch reefs are small Upper Eifelian build?ups within the fine?grained argillaceous limestone of the Hanonet Formation. Whereas the reefs themselves are not well exposed, their fossil assemblage is accessible in the hills near the town of Wellin, approximately 40xA0km SE of Dinant in Belgium. It is especially rich in massive stromatoporoids, heliolitids and other tabulate corals. They exhibit predominantly domical and bulbous morphologies. This paper focuses primarily on the palaeoautoecology of the heliolitid corals and their relationships with other organisms. Cases of mutual overgrowth between heliolitids, other corals and stromatoporids suggest a high degree of competition for space on the reefs, possibly related to the scarcity of hard substrates. Coral and stromatoporoid growth forms, as well as the prevalence of micritic matrix, point to a relatively low energy environment. However, abundant growth interruption surfaces, sediment intercalations and rejuvenations of corals suggest episodically increased hydrodynamic regime and sediment supply. It is inferred that the patch reefs developed in a relatively shallow environment, where the reefal assemblage was regularly affected by storms. Heliolitids exhibited high sediment tolerance and relied on passive sediment removal for survival. They also could regenerate effectively and commonly overgrew their epibionts, after the colony’s growth was hampered by the sediment. This is recorded in extremely abundant growth interruption surfaces, which allow the analysis of the impact of sediment influxes on the heliolitid corals. ? 2021 Lethaia Foundation. Published by John Wiley & Sons Lt

Devonian pearls and ammonoid-endoparasite co-evolution

Raised shell projections on the inner shell walls that form pits on the internal moulds of Devonian ammonoids have been known for several decades. New specimens from Morocco reveal novel details of these structures; most, if not all, of which consist of a capsule of ammonoid shell that covers tiny tubes attached to the outer (=lateral or ventral) shell wall from the inside. In accordance with comparable Recent occurrences of similar structures in molluscs, we use the term “pearls” for these structures and the pits they form on the internal moulds. The nature of these encapsulated tubes is described and discussed. Because of the presence of these tubes inside the pearls, pearl arrangement, and their similarity to Recent mollusc occurrences, the tubes are interpreted as traces of parasitoses. The pearls and pits were grouped into five types based on differences in morphology, size, and arrangement. Then, having used these traits to perform a simple cladistic analysis, the resulting cladogram was compared to the phylogeny of ammonoids. Based on this comparison, it appears likely that the parasites underwent a co-evolution with the ammonoids, which lasted 10 to 15 Ma. Patterns of evolutionary events include co-speciation, “drowning on arrival” (end of parasite lineage near base of a new host clade), and “missing the boat” (parasite lineage does not adapt to a new host clade, thus not evolving a new parasite clade). Because of the lack of fossilised soft tissue, only speculations can be made about the systematic affiliation of the parasites, their life-cycle, infection strategy, and ecological framework. Some co-occurring bivalves also have pits reminiscent to structures caused by trematodes in Recent forms. Based on the available information, the tubes are interpreted as artefacts of trematode infestations, which, if correct, would extend the fossil record of parasitic trematodes into the Early Devonian