Phylogenetic analysis of extreme morphologies: deep-sea holasteroid echinoids

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Abstract
Because of their ‘bizarre’ features, some echinoids can be considered laboratories in which to analyse the origin of extreme morphologies. The holasteroid family Urechinidae Duncan, 1889 is such a group. It is composed of 18 previously named living species in three genera: Urechinus A. Agassiz, 1879, Plexechinus A. Agassiz, 1898, and Pilematechinus A. Agassiz, 1904. With Stereopneustes, the most closely related living taxon, and the calymnid-plus-pourtalesiid clade as outgroups, a phylogenetic analysis on 35 binary characters is produced. We focus on two major aspects of morphology: test features (apical system, basicoronals, interambulacrum 5, test shape, ambulacra) and external appendages (pedicellariae, spines, sphaeridia). New data on plate architecture, morphology, and appendages are illustrated, and the position of the fossil, Chelonechinus, is discussed. We show that the Urechinidae is paraphyletic: the genus Plexechinus shares more recent common ancestry with the calymnids and pourtalesiids than with other urechinids. In order to retain the well-circumscribed clade, Pilematechinus A. Agassiz, 1904, the phylogenetic classification also recognizes two additional genera: Cystechinus A. Agassiz, 1879, and Antrechinus new genus. A key to the species is provided. The production of extreme morphologies, such as internal parental care, is explored by mapping test size, the occurrence of fascioles, and periproct position onto the phylogeny. By adding plates early in ontogeny, Pilematechinus develops radically different plate architectures from another genus of large forms, Cystechinus. In contrast, the latter gets large by exaggerating allometric trends seen in other holasteroids. Paedomorphosis and miniaturization have evolved independently in Antrechinus and some Plexechinus, but always by truncation of allometric trajectories.