Anatomy of the coelomic system in Novocrania anomala (Brachiopoda, Craniiformea) and relationships within brachiopods

Abstract

Brachiopoda is a relict phylum of marine benthic animals that have not been adequately studied with modern microscopy methods. Microscopic study may provide useful information on the evolution of the brachiopod body plan and brachiopod phylogeny. Understanding the organisation of the coelomic system is important because of its role in body form and compartmentalisation. Most brachiopods are considered to have a bipartite coelomic system; the only known exception is Lingulida, which have a tripartite coelomic system. In the present study, we provide the first complete 3D reconstruction of the coelomic system in the craniide brachiopod Novocrania anomala (Müller, 1776). Its coelomic system consists of the following five main parts, which are entirely separated from each other: 1) a pair of large brachial canals; 2) a complex system of paired small brachial canals and a perioesophageal coelom; 3) frontal coelomic chambers; 4) a main trunk coelom, which includes several semi-detached muscular chambers and mantle sinuses; and 5) a pair of posterior adductors chambers. These results indicate that the coelomic system of N. anomala (and perhaps of other craniides) is complex and cannot be considered to be bipartite or tripartite. The frontmost part of the coelomic system is represented by a pair of frontal chambers, which are considered to be a part of the lophophore but which are derived from dorsal mantle fold extensions and thus may be a part of the trunk coelomic system. A number of similarities were discovered between craniiformean and rhynchonelliformean coelomic systems, including the prominent dorsal projections of the large brachial canals and the morphological features of the perioesophageal coelom. The complex subdivision of the N. anomala trunk coelom is explained by the location and function of muscles, and by the location of several mesenteries.