Middle Devonian brachiopod-hosted sclerobiont assemblage from the northern shelf of Gondwana (Mader Basin, Morocco): Diversity, colonization patterns and relation to coeval palaeocommunities

Middle Devonian (Eifelian/Givetian transition) brachiopod-hosted sclerobiont assemblage from Gondwana (Morocco, eastern Anti-Atlas, Mader Basin) has been studied for the first time. The analysis of hundreds of brachiopod shells revealed at least 26 sclerobiont taxa, making the studied palaeoecommunity one of the most diverse Middle Devonian sclerobiont assemblage reported to date. The palaeocommunity is dominated by encrusting organisms, in particular bryozoans, foraminifers, ascodictyids and microconchids, whereas bioeroders are represented by Clionolithes and Oichnus ichnospecies. As supported by microfacies, fossil preservation and the presence of encrusting Rothpletzella algae, the sclerobionts developed in a calm, euphotic, offshore marine environment situated below normal wave base. Prevalence of articulated shells without signs of either extensive mineral staining, or taphonomic alteration resulting from hydrodynamic processes, as well as preservation of some delicate encrusters (Vinella, Rothpletzella), indicate that the palaeocommunity did not undergo extensive time-averaging. The hosts were colonized syn vivo, as evidenced by the epibiont growth near the commissure margin, the presence of shell malformations and even embedded auloporid corals and cornulitid tubeworms. Competition for space among sclerobionts was apparently limited, as suggested by the very low number of overgrowth interactions, indicating that the shelly substrate available for colonization was abundant. The presence of the same dominant groups of sclerobionts on different host taxa indicates a lack of species-selectivity by these colonizing sclerobionts. It also suggests that the sclerobiont species pool in the environment was generally similar throughout the deposition of the sampled rock interval. As testified by the host brachiopod and bryozoan species, the sclerobiont assemblage from this locality of northern Gondwana had strong affinities with those from the southern margin of Laurussia, most likely reflecting strong connectivity between these regions provided by the South Equatorial – North Gondwana current system during the late Eifelian Kačák transgressive episode.