A primitive starfish ancestor from the Early Ordovician of Morocco reveals the origin of crown group Echinodermata

The somasteroids are Ordovician star-shaped animals widely regarded as ancestors of Asterozoa, the group of extant echinoderms that includes brittle stars and starfish. The phylogenetic position of somasteroids makes them critical for understanding the origin and early evolution of crown group Echinodermata. However, the early evolution of asterozoans, the origin of their distinctive body organization and their relationships with other Cambrian and Ordovician echinoderms, such as edrioasteroids, blastozoans, crinoids, and other asterozoans, remain problematic due to the difficulties of comparing the calcitic endoskeleton of these disparate groups. Here we describe the new somasteroid Cantabrigiaster fezouataensis from the Early Ordovician (Tremadocian) Fezouata Lagerstätte in Morocco. Cantabrigiaster shares with other somasteroids the presence of rod-like virgal ossicles that articulate with the ambulacrals, but differs from all other known asterozoans in the absence of adambulacral ossicles defining the arm margins. The unique arm construction evokes parallels with non-asterozoan echinoderms. Developmentally informed Bayesian and parsimony based phylogenetic analyses, which reflect the homology of the biserial ambulacral ossicles in Paleozoic echinoderms according to the Extraxial-Axial Theory, recover Cantabrigiaster as basal within stem group Asterozoa. Our results indicate that Cantabrigiaster is the earliest diverging stem group asterozoan, revealing the ancestral morphology of this major clade and clarifying the affinities of problematic Ordovician taxa. Somasteroids are resolved as a paraphyletic grade within stem and crown group Asterozoa (starfishes), whereas stenuroids are paraphyletic within stem group Ophiuroidea (brittle stars). Cantabrigiaster also illuminates the relationship between Ordovician crown group Echinodermata and its Cambrian stem lineage, which includes sessile forms with incipient radial symmetry such as edrioasteroids and blastozoans. The contentious Pelmatozoa hypothesis (i.e. monophyly of blastozoans and crinoids) is not supported; instead, blastozoans represent the most likely sister-taxon of crown group Echinodermata. Author summary Starfish and brittle stars, collectively known as asterozoans, constitute a diverse and ecologically successful group of echinoderms that first appear in the fossil record some 480Ma. However, the early evolution of asterozoans, the origin of their distinctive body organization, and their phylogenetic relationships with Cambrian echinoderms remain largely unresolved. We describe Cantabrigiaster fezouataensis gen. et sp. nov., a primitive asterozoan from the Fezouata Lagerstätte, Morocco, with a unique endoskeletal arm organization that reveals the ancestral morphology of this major clade. Bayesian and parsimony based phylogenetic analyses indicate that Cantabrigiaster is the earliest diverging stem group asterozoan, and resolve the phylogenetic position of Ordovician asterozoans such as somasteroids. Our analyses clarify the origin of crown group echinoderms relative to their problematic Cambrian stem group representatives.