818


Journal of Paleontology 91(4):815–828


Table 2. Genera, species, and coding references used in the preliminary phylogenetic analysis. Asterisk (*) denotes non–type species coded due to poor preservation of the type. First appearance is given for genus.


Genus and species coded


Blastozoa Cheirocystis fultonensis* Sumrall and Schumacher, 2002 Echinosphaerites aurantium (Gyllenhaal, 1772) Eumorphocystis multiporata Branson and Peck, 1940 Glyptocystella loeblichi (Bassler, 1943) Protocrinites rugatus* Bockelie, 1984 Rhopalocystis destombesi Ubaghs, 1963


Cladida Aethocrinus moorei Ubaghs, 1969


Apektocrinus ubaghsi Guensburg and Sprinkle, 2009 Compagicrinus fenestratus Jobson and Paul, 1979 Elpasocrinus radiatus Sprinkle and Wahlman, 1994


Camerata, Diplobathrida Archaeocrinus microbasalis* (Billings, 1857)


Cnemecrinus fillmorensis Guensburg and Sprinkle, 2003 Euptychocrinus skopaios* Brower, 1994 Neoarchaeocrinus pyriformis (Billings, 1857)


Proexenocrinus inyoensis Strimple and McGinnis, 1972 Quechuacrinus tisca Guensburg and Waisfeld, 2015 Reteocrinus alveolatus* (Miller and Gurley, 1894) Rhaphanocrinus subnodosus (Walcott, 1883)


Trichinocrinus terranovicus Moore and Laudon, 1943b


Camerata, Monobathrida Abludoglyptocrinus charltoni (Kolata, 1975) Celtocrinus ubaghsi Donovan and Cope, 1989


Eknomocrinus wahwahensis Guensburg and Sprinkle, 2003 Glyptocrinus decadactylus Hall, 1847 Xenocrinus baeri* (Meek, 1872)


Protocrinoidea Glenocrinus globularis Guensburg and Sprinkle, 2003 Titanocrinus sumralli Guensburg and Sprinkle, 2003


Tremadocian Tremadocian Floian Floian


Darriwilian Tremadocian Katian


Darriwilian Floian Floian


Sandbian


Darriwilian Darriwilian


Sandbian Floian


Tremadocian Sandbian Katian


Tremadocian Tremadocian


and the Birmingham Lapworth Museum. Coding of specimens was supplemented using primary taxonomic literature (Table 3). In instances where museum specimens were inaccessible, taxa were coded using primary taxonomic literature. For each genus, the species selected for coding was typically the type species except in cases where a non–type species preserved more characters than the type or where the type species was post- Ordovician in age. Species were coded for 112 discrete char- acters, 64 binary and 48 multistate, that were selected to capture morphological variation among Ordovician camerates and are inferred to represent homologous structures among sampled taxa (Foote, 1994, 1999; Ausich, 1996; Kammer et al., 2013; Wright, 2015) (Supplemental Data 2). Of the characters coded, 85 proved to be parsimony-informative. The Tremadocian diplobathrid Eknomocrinus was designated as the outgroup based on the results of the preliminary analysis that identified Cnemecrinus and Eknomocrinus as sister to the Crinoidea. Although both taxa are Tremadocian in age, Eknomocrinus is from a lower stratigraphic horizon than Cnemecrinus (Guensburg and Sprinkle, 2003). Designation of Cnemecrinus instead of Eknomocrinus as the outgroup, however, resulted in only trivial changes in ingroup topology.


Phylogenetic analysis.—To assess evolutionary relationships among the Ordovician Camerata, a parsimony analysis was con- ducted in PAUP* v. 4.0a147 (Swofford, 2003) using a heuristic search with 1,000 random addition sequences. TBR was used for the branch-swapping algorithm with no reconnection limit, and branches with a maximum length of zero were collapsed. All characters were left unordered and equally weighted. The parsimony analysis recovered 801 most parsimonious trees with lengths of 634 steps per tree. Strict consensus of the 801 most


Ubaghs, 1969; Rozhnov, 1988 Guensburg and Sprinkle, 2009


Rozhnov, 1988; Webster and Maples, 2006 Sprinkle and Wahlman, 1994


Wachsmuth and Springer, 1897 Guensburg and Sprinkle, 2003 Brower, 1994


Billings, 1859; Wachsmuth and Springer, 1897 Strimple and McGinnis, 1972; Ausich, 1985 Guensburg and Waisfeld, 2015 Springer, 1911; Guensburg, 1984 Wachsmuth and Springer, 1897


Moore and Laudon, 1943b; Ausich, Bolton, and Cummings, 1998 Kolata, 1975 Donovan and Cope, 1989


Guensburg and Sprinkle, 2003 Wachsmuth and Springer, 1897


Wachsmuth and Springer, 1897; Brower, 1974


Guensburg and Sprinkle, 2003 Guensburg and Sprinkle, 2003


parsimonious trees resulted in a tree topology with little resolution (Supplemental Data 3). Subsequently, the characters were reweighted using the rescaled consistency (RC) function in PAUP*. This method of reweighting adjusts the weight of characters in an attempt to minimize homoplasy in the analysis (Farris, 1989; Smith, 1994b). After reweighting using RC, the analysis was run for 1,000 random addition sequences using the same search parameters as the initial search, and a single most parsimonious tree was recovered (Fig. 2). For the reweighted analysis, CI and RI values were recorded for the recovered tree, and bootstrap values and Bremer support were calculated using PAUP*.


Additional analyses.—Additional character analyses were conducted in Mesquite (Maddison and Maddison, 2015) for the single most parsimonious tree recovered from the reweighted analysis (Fig. 2). Using the R package STRAP (Bell and Lloyd, 2015), the most parsimonious tree was plotted against the observed stratigraphic ranges of sampled genera to produce a time-scaled phylogeny (Fig. 3). Stratigraphic ranges of genera used to time-scale the tree were tabulated from Webster (2003) and updated to include new taxa and match current divisions of the International Chronostratigraphic Chart. Stratigraphic congruence metrics were also calculated for


the tree using STRAP to assess the fit of the recovered tree to observed stratigraphic ranges. The stratigraphic congruence measures used were the stratigraphic consistency index (SCI; Huelsenbeck, 1994), relative completeness index (RCI; Benton and Storrs, 1994), the gap excess ratio (GER; Willis, 1999), and Manhattan stratigraphic measure* (MSM*; Siddall, 1998; Pol and Norell, 2001). For each measure, the calculated value was compared to a null model of 1,000 trees generated through


First appearance


Katian Katian


Sandbian Sandbian Sandbian


Ordovician Coding references


Sumrall and Schumacher, 2002 Bockelie, 1981, 1984 Branson and Peck, 1940 Sprinkle, 1982a Bockelie, 1984 Ubaghs, 1963


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