Journal of Paleontology, 91(4), 2017, p. 829–846 Copyright © 2017, The Paleontological Society 0022-3360/17/0088-0906 doi: 10.1017/jpa.2016.142
Phylogenetic taxonomy and classification of the Crinoidea (Echinodermata)
David F. Wright,1,* William I. Ausich,1 Selina R. Cole,1 Mark E. Peter,1 and Elizabeth C. Rhenberg2 1School of Earth Sciences, 125 South OvalMall, Ohio State University, Columbus, OH 43210, USA ⟨
wright.1433@osu.edu⟩, ⟨
ausich.1@osu.edu⟩, ⟨
cole.678@osu.edu⟩, ⟨
peter.1@osu.edu⟩
2Department of Geology, Earlham College, 801 National Road West, Richmond, IN 47374, USA ⟨
rhenbel@earlham.edu⟩
Abstract.—A major goal of biological classification is to provide a system that conveys phylogenetic relationships while facilitating lucid communication among researchers. Phylogenetic taxonomy is a useful framework for defining clades and delineating their taxonomic content according to well-supported phylogenetic hypotheses. The Crinoidea (Echinodermata) is one of the five major clades of living echinoderms and has a rich fossil record spanning nearly a half billion years. Using principles of phylogenetic taxonomy and recent phylogenetic analyses, we provide the first phylogeny-based definition for the Clade Crinoidea and its constituent subclades. A series of stem- and node-based defi- nitions are provided for all major taxa traditionally recognized within the Crinoidea, including the Camerata, Disparida, Hybocrinida, Cladida, Flexibilia, and Articulata. Following recommendations proposed in recent revisions, we recog- nize several new clades, including the Eucamerata Cole 2017, Porocrinoidea Wright 2017, and Eucladida Wright 2017. In addition, recent phylogenetic analyses support the resurrection of two names previously abandoned in the crinoid taxonomic literature: the Pentacrinoidea Jaekel, 1918 and Inadunata Wachsmuth and Springer, 1885. Last, a phyloge- netic perspective is used to inform a comprehensive revision of the traditional rank-based classification. Although an attempt was made to minimize changes to the rank-based system, numerous changes were necessary in some cases to achieve monophyly. These phylogeny-based classifications provide a useful template for paleontologists, biologists, and non-experts alike to better explore evolutionary patterns and processes with fossil and living crinoids.
Introduction
Crinoids are a diverse, long-lived clade of echinoderms with a fossil record spanning nearly half a billion years and are repre- sented by more than 600 species living in marine ecosystems today (Hess et al., 1999). The geologic history of crinoids is revealed through a highly complete, well-sampled fossil record (Foote and Raup, 1996; Foote and Sepkoski, 1999) displaying a complex pageant of evolutionary radiation, extinction, ecologic innovation, and morphologic diversification (Ausich and Bottjer, 1982; Ausich et al., 1994; Foote, 1999; Peters and Ausich, 2008; Deline and Ausich, 2011; Gorzelak et al., 2015). The spectacular fossil record of crinoids is greatly enriched and complemented by detailed biologic studies on living species. These studies facilitate opportunities to synthesize information from fossil and extant forms. For example, comparative studies between fossil and living crinoid species have provided insight into species ecology and niche dynamics (Meyer and Macurda, 1977; Ausich, 1980; Roux, 1987; Kitazawa et al., 2007; Baumiller, 2008), established developmental bases for mor- phologic homologies (Shibata et al., 2015a), and informed phylogenetic hypotheses (Simms and Sevastopulo, 1993; Rouse et al., 2013). Thus, crinoids form a data-rich model system for exploring major questions in the history of life.
* Present address: Department of Paleobiology, National Museum of Natural History, The Smithsonian Institution, P.O. Box 37012, MRC 121, Washington, DC 20013-7012, USA ⟨
wrightda@si.edu⟩
Given their general significance and broad scientific utility
across multiple disciplines of inquiry, it is paramount that the biological classification of crinoids reflects their evolutionary heritage. Numerous emendations and informal suggestions for major taxonomic revisions have been opined over the past few decades (e.g., Kelly, 1986; Simms and Sevastopulo, 1993; Ausich, 1998a, 1998b; Webster and Jell, 1999; Hess and Messing, 2011), but the most recent comprehensive revision to crinoid classification is the 1978 Treatise on Invertebrate Paleontology (Moore and Teichert, 1978). Since publication of the Treatise, the value of revising rank-based systematic classifications to be consistent with phylogenetic hypotheses and/or the explicit use of phylogenetic taxonomy (sensu de Quieroz and Gauthier, 1990; Sereno, 1999, 2005) has become increasingly common in paleontology (e.g., Smith, 1984, 1994; Holtz, 1996, 1998; Sereno, 1997; Padian et al., 1999; Brochu and Sumrall, 2001; Carlson, 2001; Carlson and Leighton, 2001; Brochu, 2003; Forey et al., 2004; Sereno et al., 2005; Butler et al., 2008; Kelley et al., 2013). We agree with these authors that all named taxa in a biological classification system should ideally represent clades (i.e., monophyletic groups). The development of phylogeny-based classifications is not without difficulties or criticism (e.g., Benton, 2000, 2007). However,we advocate that recent advances in understanding the phylogenetic relationships of major crinoid lineages make the biological classification of the Crinoidea ripe for revision. Agreat strength of so-called ‘phylogenetic taxonomy’ is its potential for increasing nomenclatural stability (de Quieroz and
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