Myoanatomy of the Lophophore in Adult Phoronids and the Evolution of the Phoronid Lophophore.

Confocal laser scanning microscopy was used to study the myoanatomy of the lophophore of three phoronids with different types of lophophore: Phoronis ijimai, Phoronis australis, and Phoronopsis harmeri. A four-part ground plan of the lophophoral musculature was detected in all three species and was previously reported for Phoronis ovalis. The ground plan includes (i) a circular muscle, (ii) longitudinal muscles of the tentacular lamina, (iii) groups of paired distal muscles of the tentacular lamina, and (iv) frontal and abfrontal muscles of the tentacles. In P. australis, the tentacular lamina contains strong abfrontal and numerous frontal muscles. Phoronis harmeri has an inner circular muscle and arch-like muscles. Among all studied phoronids, the four-part ground plan of the lophophoral musculature is least complex in P. ijimai, which has a horseshoe-shaped lophophore. The results suggest two possible scenarios by which the morphology of the phoronid lophophore has transformed over evolutionary time. According to the first scenario, the morphology of the ancestral horseshoe-shaped lophophore became more complicated in the case of most phoronids but became simplified in the case of P. ovalis and bryozoans. According to the second scenario, the lophophore gradually transformed from a simple oval shape to a horseshoe shape and then to a spiral shape. The four-part ground plan of the lophophoral musculature is also present in bryozoans, which is consistent with the view that the lophophorates are monophyletic.