Patterns of cornulitid encrustation on the Late Devonian brachiopod shells from Russia

Upper Devonian brachiopods from the Central Devonian Field, Russia have been investigated with respect to encrustation patterns of cornulitids. These Palaeozoic encrusters were very characteristic component of shelly substrates during the Devonian and may serve as valuable objects for palaeoecological studies, especially those concerning the encrusters-host relationships. On the basis of rich material, it has been shown that cornulitid abundance significantly increased in the lower Famennian. Their disparity also increased from one morphotype (robust) present in the upper Frasnian to two distinct morphotypes (robust and slender, presumably representing two different species) in the lower Famennian. The analysis of the spatial occurrence of cornulitids on two dominant lower Famennian brachiopod species (Cyrtospirifer zadonicus and Ripidiorhynchus huotinus) showed, that these encrusters preferred the marginal parts of the shells. This, together with a dominant directional growth of cornulitids toward the commissure, indicates that cornulitids benefited from food brought by feeding currents produced by the brachiopod's lophophore. The unequivocal evidence for syn vivo association of the cornulitids and their brachiopod hosts is the presence of distinct shell malformations caused by cornulitid growth affecting the host's shell-secreting epithelium, preserved on some lower Famennian specimens. Interestingly, these malformations were caused by a slender cornulitid morphotype only. Thus, it is evident that during food gathering from the inhalant currents, this cornulitid species exerted a distinct, negative effect on the brachiopod - a deviation of its shell growth. In this case, we may consider this particular cornulitid species as ectoparasite rather than commensal with respect to its host.