Phylogeny and classification of the Asteroidea (Echinodermata)

Post-Palaeozoic asteroids share a large number of derived characters of the ambulacral column and the mouth frame, and constitute the crown group of the monophyletic group Asteroidea. This crown group is here called the Neoasteroidea (new subclass). The stem species of the crown group lived in the Permian or early Triassic and so the evolution of the asteroids parallels that of the echinoids. Character distribution within the Neoasteroidea, especially morphology of the skeleton, digestive system, larvae and tube feet, allows subdivision into four orders (Paxillosida, Notomyotida, Valvatida, Forcipulatida). The latter three orders possess the synapomorphy of suckered tube feet and are united as the Surculifera (new superorder); the Paxillosida are their primitive sister group. Palaeozoic asteroids represent the stem group of the class, and may be divided into plesions according to the order of appearance of synapomorphies with the crown group. Classification of Palaeozoic asteroids requires much further study. The appearance of new characters within the crown group asteroids, such as suckered tube feet, implies that these were absent in the stem group. The range of life-habits possible in Palaeozoic asteroids can thus be partly deduced from evidence provided by living asteroids. Palaeozoic asteroids are deduced to have lacked suckered tube feet and were presumably unable to evert the stomach; hence they were precluded from life on hard substrates and extraoral feeding on epifaunal organisms. It is suggested that they lived on soft substrates by deposit feeding, scavenging and predation on small benthos.