THALIACEANS, THE NEGLECTED PELAGIC RELATIVES OF ASCIDIANS: A DEVELOPMENTAL AND EVOLUTIONARY ENIGMA.

Most developmental biologists equate tunicates to the sessile ascidians, including Ciona intestinalis, and the pelagic appendicularians, in particular Oikopleura dioica. However, there exists a third group of tunicates with a pelagic lifestyle, the thaliaceans, which include salps, pyrosomes, and doliolids. Although thaliaceans have raised the curiosity offamous zoologists since the 18th century, the difficulty of observing and experimentally manipulating them has led to many controversies and speculations about their life cycles and developmental strategies, the phylogenetic relationship within the group and with other tunicates, and the drivers of speciation in these widely distributed animals living in a seemingly uniform environment. Here, we take a historical perspective to summarize 250 years of work on this intriguing group of animals, and explore how modern genomics and imaging approaches are starting to solve fascinating evolutionary and developmental riddles. Recent molecular analyses support previous morphological evidence that ascidians are not monophyletic and that thaliaceans evolved from a sessile ascidian-like ancestor. In parallel, preliminary live-imaging and gene-expression data offer exciting entry points to understand how the adoption of a pelagic lifestyle led to drastic modifications in the morphology, embryology, and life cycle of these tunicates, compared to their sessile ancestor.