The bulbus arteriosus of the holocephalan heart: gross anatomy, histomorphology, pigmentation, and evolutionary significance.

This study was designed to determine whether the outflow tract of the holocephalan heart is composed of a myocardial conus arteriosus and a non-myocardial bulbus arteriosus, as is the case in elasmobranchs. This is a key issue to verify the hypothesis that these two anatomical components existed from the onset of the jawed vertebrate radiation. The Holocephali are the sister group of the elasmobranchs, sharing with them a common, still unknown Palaeozoic ancestor. The sample examined herein consisted of hearts from individuals of four species, two of them belonging to the Chimaeridae and the other two to the Rhinochimaeridae. In all specimens, the cardiac outflow tract consisted of a conus arteriosus, with myocardium in its walls and two rows of valves at its luminal side, and an intrapericardial bulbus arteriosus shorter than the conus and devoid of valves. The bulbus, mainly composed of elastin and smooth musculature, was covered by the epicardium and crossed longitudinally by coronary artery trunks. These findings give added support to the viewpoint that the outflow tract of the primitive heart of the gnathostomes was not composed of a single component, but two, the conus and the bulbus. All rabbitfish (Chimaera monstrosa) examined had pigment cells over the surface of the heart. The degree of pigmentation, which varied widely between individuals, was particularly intense in the cardiac outflow tract. Pigment cells also occurred in the bulbus arteriosus of one of the two hearts of the straightnose rabbitfish (Rhinochimaera atlantica) included in the study. The cells containing pigment, presumably derived from the neural crest, were located in the subepicardium.