Evidence for Polyphyly of the Genus <i>Scrupocellaria</i> (Bryozoa: Candidae) Based on a Phylogenetic Analysis of Morphological Characters

<div><p>The bryozoan genus <i>Scrupocellaria</i> comprises about 80 species in the family Candidae. We propose a hypothesis for the phylogenetic relationships among species assigned to <i>Scrupocellaria</i> to serve as framework for a phylogenetic classification using 35 morphological characters. Our results suggest that the genus <i>Scrupocellaria</i> is polyphyletic. <i>Scrupocellaria s. str.</i> is redefined according to four morphological features: vibracular chamber with a curved setal groove, ooecium with a single ectooecial fenestra, two axillary vibracula, and a membranous operculum with a distinct distal rim. Thus, the genus includes only 11 species: <i>Scrupocellaria aegeensis</i>, <i>Scrupocellaria delilii</i>, <i>Scrupocellaria harmeri</i>, <i>Scrupocellaria incurvata</i>, <i>Scrupocellaria inermis</i>, <i>Scrupocellaria intermedia</i>, <i>Scrupocellaria jullieni</i>, <i>Scrupocellaria minuta</i>, <i>Scrupocellaria puelcha</i>, <i>Scrupocellaria scrupea</i>, and <i>Scrupocellaria scruposa</i>. The monophyly of <i>Cradoscrupocellaria</i> is supported and five new genera are erected: <i>Aquiloniella</i> n. gen., <i>Aspiscellaria</i> n. gen., <i>Paralicornia</i> n. gen., <i>Pomocellaria</i> n. gen. and <i>Scrupocaberea</i> n. gen. Two other new genera, <i>Bathycellaria</i> n. gen. and <i>Sinocellaria</i> n. gen., are erected to accommodate two poorly known species, <i>Scrupocellaria profundis</i> Osburn and <i>Scrupocellaria uniseriata</i> Liu, respectively. <i>Scrupocellaria congesta</i> is tentatively assigned to <i>Tricellaria</i>. Fifteen species are reassigned to <i>Licornia</i>: <i>Licornia cookie</i> n. comb., <i>Licornia micheli</i> n. comb., <i>Licornia milneri</i> n. comb., <i>Licornia curvata</i> n. comb., <i>Licornia diegensis</i> n. comb., <i>Licornia drachi</i> n. comb., <i>Licornia mexicana</i> n. comb., <i>Licornia pugnax</i> n. comb., <i>Licornia raigadensis</i> n. comb., <i>Licornia regularis</i> n. comb., <i>Licornia resseri</i> n. comb., <i>Licornia securifera</i> n. comb., <i>Licornia spinigera</i> n. comb., <i>Licornia tridentata</i> n. comb., and <i>Licornia wasinensis</i> n. comb. <i>Notoplites americanus</i> n. name is proposed as a replacement name for <i>Scrupocellaria clausa</i> Canu & Bassler. Three fossil species are reassigned to <i>Canda</i>: <i>Canda rathbuni</i> n. comb., <i>Canda triangulata</i> n. comb. and <i>Canda williardi</i> n. comb. A species is reassigned to <i>Notoplites</i>, <i>Notoplites elegantissima</i> n. comb. The generic assignment of eleven species of <i>Scrupocellaria</i>, including <i>Scrupocellaria macandrei</i>, remains uncertain.</p></div>