A phylogeny of the hemichordates based on morphological characters

Abstract

A comprehensive review of literature on all 15 genera constituting the phylum Hemichordata resulted in a morphological matrix of 105 characters. The echinoderms, tunicates, cephalochordates, and vertebrates were included in the analysis, and the cnidarians, polychaetes, and sipunculids were employed as outgroup taxa. The consensus tree supported the traditional view of a monophyletic Hemichordata, Echinodermata, Ambulacraria, and Chordata. The enteropneust families Spengelidae and Ptychoderidae were each monophyletic and sister-taxa, but there was no resolution among the family Harrimaniidae. A detailed sensitivity analysis provided (i) tree lengths of competing evolutionary hypothesis and (ii) a test of monophyly of groups under a variety of evolutionary models. It is argued that the ancestral deuterostome was a benthic vermiform organism with a terminal mouth and anus, well-developed circular and longitudinal muscles, a simple nerve plexus with little sign of regionalization, a pharynx with gill slits and collagenous gill bars, a cluster of vacuolated cells with myofilaments, produced iodotyrosine, and displayed direct development. The pterobranchs have lost many of these features as a consequence of evolving a small body size and living in tubes, but these features exist in present-day enteropneusts, suggesting that they are a plausible model for the proximate ancestor of deuterostomes.