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Generic concepts in the Actinocrinitidae Austin and Austin, 1842 (class Crinoidea) and evaluation of generic assignments of species

Elizabeth C. Rhenberg, William I. Ausich and Thomas W. Kammer

Journal of Paleontology / Volume 89 / Issue 01 / January 2015, pp 1 - 19 DOI: 10.1017/jpa.2014.2, Published online: 09 March 2015

Link to this article: http://journals.cambridge.org/abstract_S002233601400002X

How to cite this article: Elizabeth C. Rhenberg, William I. Ausich and Thomas W. Kammer (2015). Generic concepts in the Actinocrinitidae Austin and Austin, 1842 (class Crinoidea) and evaluation of generic assignments of species. Journal of Paleontology, 89, pp 1-19 doi:10.1017/jpa.2014.2

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Downloaded from http://journals.cambridge.org/JPA, IP address: 157.182.154.151 on 10 Mar 2015 Journal of Paleontology, 89(1), 2015, p. 1–19 Copyright © 2015, The Paleontological Society 0022-3360/15/0088-0906 doi: 10.1017/jpa.2014.2

Generic concepts in the Actinocrinitidae Austin and Austin, 1842 (class Crinoidea) and evaluation of generic assignments of species

Elizabeth C. Rhenberg1, William I. Ausich1, and Thomas W. Kammer2

1 Geology Department, Earlham College, Richmond, VA 47374 〈[email protected]〉 2Department of Geology and Geography, West Virginia University, Morgantown, WV 26506-6300 〈[email protected]〉; 〈[email protected]〉

Abstract.—The family Actinocrinitidae was a significant contributor to the global biodiversity peak of crinoids that occur- red during the Mississippian and is referred to as the “Age of Crinoids.” Although the actinocrinitids are a major component of that high diversity, they are also a source of much taxonomic confusion. Previously, generic concepts were not applied equally between Europe and North America creating disparity in the definition of genera. In this contribution, all genera are defined objectively by discrete characters, and the generic assignments of all species are reevaluated. Twenty genera are described. A total of 206 species were evaluated of which 56 species and one taxon in open nomenclature are reassigned to different genera, 21 species are designated as nomina dubia, and three species and one genus are now incertae sedis. A phylogenetic hypothesis is presented for the relationships of the genera of Actinocrinitidae genera based on a parsimony-based analysis and plotted against stratigraphic ranges. Although groupings were revealed in this analysis, the Actinocrinitidae cannot be readily subdivided into subfamilies. Rapid diversification occurred during the Tournaisian following the Hangenberg Extinction of probable fish predators. The late Devonian (Famennian) occurrence of the highly derived genera Abactinocrinus and Physetocrinus suggests there is a more extensive, but undocumented, evolutionary history for the Actinocrinitidae during the Devonian.

Introduction been applied as species-level diagnostic characters among North American forms. For example, ray grouping and lobation helps The Actinocrinitidae is an important family of Mississippian cri- to define European genera, but this has been used to differentiate noids due to their considerable genus diversity and abundance of North American species of Actinocrinites. Unfortunately, the specimens. Actinocrinitids encompass a large degree of morpho- solution for this is reassignment of many well-known North logical disparity produced by the complex arrangements of their American species of “Actinocrinites” to other genera. Although multi-plated calyxes. Because there are innumerable specimens in nomenclaturally messy, this situation is not surprising given that many Mississippian faunas and because we suspect that hybridi- many nineteenth century broadly defined genera have been zation was a complicating factor as demonstrated in Eretmocrinus subdivided into numerous new genera, such as Cyathocrinites, (Ausich and Meyer, 1994), and suggested for the Batocrinidae Phanocrinus, Platycrinites, and Poteriocrinites (see Webster, (Ausich and Kammer, 2010), this is a complex group. Both 2003). Recognition of well-defined generic concepts is a pre- Wachsmuth and Springer (1897) and Bowsher (1955) worked to requisite for a modern understanding of the phylogeny, paleo- clarify generic definitions among North American actinocrinitids, biogeography, and evolutionary paleoecology of these crinoids. but many issues still remain. First and most significant among This study is a continuation of the efforts to redefine the these problems is that the generic concept for Actinocrinites,as generic concepts of monobathrid camerates (Ausich and Kammer, typified by Actinocrinites triacontadactylus Miller, 1821, the type 2008, 2009, 2010). Revising the camerates is an important step in species of the genus, has not been applied consistently to North compiling an accurate database to evaluate their evolutionary American faunas. paleoecology of Mississippian crinoids, including an examination Ausich and Sevastopulo (2001) designated a neotype for of the global, generic relationships among longevity, eurytopy, and Actinocrinites triacontadactylus; and with this definition re- evolutionary paleoecology (Ausich and Kammer, 2010). The established, Ausich and Sevastopulo (2001) and Ausich and camerates are of particular interest because they reached their peak Kammer (2006) made generic reassignments to all actinocrinitids diversity during the Tournaisian and are one of the most diverse from Ireland, England, and Wales. Species-level systematics has groups of crinoids during this time (Kammer and Ausich, 2006; only been completed for Ivorian faunas (Ausich and Sevastopulo, Sallan et al., 2011). 2001). A total of six actinocrinitid genera are now recognized from In this contribution, the goals are to define all Actinocrinitidae Ireland, England, and Wales. genera using objective characters and to assign all species to the The primary source of confusion regarding the systematics correct genus. This study treated 206 species, with 57 species of North American (and other) actinocrinitids is that genus-level (and one species left in open nomenclature) reassigned to different diagnostic characters that define European actinocrinitids have genera (Table 1); 20 species were designated as nomina dubia;and 1 Journal of Paleontology 89(1):1–19 2

Table 1. Listing of actinocrinitid species that have a revised generic assignment

Previous assignment (Webster, 2003) New assignment ?Actinocrinites polydactylus Miller, 1821 Nomen dubium Aacocrinus acylus Webster and Jell, 1999 Nomen dubium Aacocrinus algerianensis Webster et al., 2004 Thinocrinus algerianensis (Webster et al., 2004) Aacocrinus enigmaticus Webster and Lane, 1897 Thinocrinus enigmaticus (Webster and Lane, 1897) Aacocrinus triarmatus Brower, 1967 Nomen dubium Actinocrinites alatus Wright, 1955a Thinocrinus alatus (Wright, 1955a) Actinocrinites anchorensis Webster and Lane, 1987 Thinocrinus anchorensis (Webster and Lane, 1987) Actinocrinites becharensis Webster et al., 2004 Nunnacrinus becharensis (Webster et al., 2004) Actinocrinites benedicti (Miller, 1892) Thinocrinus benedicti (Miller, 1892) Actinocrinites blairi (Miller and Gurley, 1897) Thinocrinus blairi (Miller and Gurley, 1897) Actinocrinites botruosus (Miller and Gurley, 1895) Thinocrinus botruosus (Miller and Gurley, 1895) Actinocrinites brevispina (Wanner, 1924) Thinocrinus brevispina (Wanner, 1924) Actinocrinites brouweri (Wanner, 1924) Thinocrinus brouweri (Wanner, 1924) Actinocrinites c.f. brouweri (Wanner, 1924) Nomen dubium Actinocrinites carinatus (Wanner, 1937) Thinocrinus carinatus (Wanner, 1937) Actinocrinites caryocrinoides (M’Coy, 1844) Nomen dubium Actinocrinites combinatus Webster et al., 2004 Thinocrinus combinatus (Webster et al., 2004) Actinocrinites constrictus (M’Coy, 1844) Nomen dubium Actinocrinites coplowensis Wright, 1955a Sampsonocrinus coplowensis (Wright, 1955a) Actinocrinites decadactylus (Gilbertson in Portlock, 1843) Nomen dubium Actinocrinites dilatus (Wanner, 1924) Thinocrinus dilatus (Wanner, 1924) Actinocrinites erraticus (Miller and Gurley, 1893) Nunnacrinus erraticus (Miller and Gurley, 1893) Actinocrinites eximius (Kirk, 1943) Thinocrinus subpulchellus (Miller and Gurley, 1896c) Actinocrinites exornatus (Wanner, 1937) Thinocrinus exornatus (Wanner, 1937) Actinocrinites gibsoni (Miller and Gurley, 1893) Thinocrinus gibsoni (Miller and Gurley, 1893) Actinocrinites goldfussi (Schmidt, 1930) Nomen dubium Actinocrinites granulatus (Wright, 1955a) Thinocrinus granulatus (Wright, 1955a) Actinocrinites granulatus Goldfuss, 1831 Nomen dubium Actinocrinites griffithi (Wachsmuth and Springer, 1897) Thinocrinus subpulchellus (Miller and Gurley, 1896c) Actinocrinites higuchisawensis (Minato, 1951) Nomen dubium Actinocrinites intermedius Wright, 1955a Iotacrinus intermedius (Wright, 1955a) Actinocrinites laevis Miller, 1821 Nomen dubium Actinocrinites lowei (Hall, 1858) Thinocrinus lowei (Hall, 1858) Actinocrinites marcoui (Collingnon, 1924) Nomen dubium Actinocrinites multiradiatus (Shumard, 1857) Thinocrinus multiradiatus (Shumard, 1857) Actinocrinites multiramosus altidorsatus (Rowley, 1904) Thinocrinus multiramosus altidorsatus (Rowley, 1904) Actinocrinites nodosus Wright, 1955a Thinocrinus nodosus (Wright, 1955a) Actinocrinites ohmoriensis (Minato, 1951) Nomen dubium Actinocrinites permicus (Wanner, 1924) Thinocrinus permicus (Wanner, 1924) Actinocrinites pernodosus (Hall, 1858) Thinocrinus pernodosus (Hall, 1858) Actinocrinites plagosus (Miller and Gurley, 1893) Nunnacrinus plagosus (Miller and Gurley, 1893) Actinocrinites probolos Ausich and Kammer, 1991 Thinocrinus probolos (Ausich and Kammer, 1991) Actinocrinites pusillus (M’Coy, 1844) Nomen dubium Actinocrinites rotundatus Wright, 1955a Thinocrinus rotundatus (Wright, 1955a) Actinocrinites scitulus (Miller and Gurley, 1897) Thinocrinus scitulus (Miller and Gurley, 1897) Actinocrinites sedaliensis (Miller and Gurley, 1895) Sampsonocrinus sedaliensis (Miller and Gurley, 1895) Actinocrinites semimultiramosus (Whitfield, 1900) Thinocrinus semimultiramosus (Whitfield, 1900) Actinocrinites skourensis (Termier, G. and Termier, H., 1950) Nomen dubium Actinocrinites sobrinus (Miller and Gurley, 1896c) Cusacrinus sobrinus (Miller and Gurley, 1896c) Actinocrinites spectabilis (Miller and Gurley, 1896c) Cusacrinus spectabilis (Miller and Gurley, 1896c) Actinocrinites spinaetectus (Wanrer, 1937) Incertae sedis Actinocrinites subpulchellus (Miller and Gurley, 1896c) Thinocrinus subpulchellus (Miller and Gurley, 1896c) Actinocrinites subscitulus (Miller and Gurley, 1896c) Cusacrinus subscitulus (Miller and Gurley, 1896c) Actinocrinites tenuis (de Koninck and Le Hon, 1854) Nomen dubium Actinocrinites tenuistriatus (Phillips, 1841) Nomen dubium Actinocrinites tricuspidatus (de Koninck and Le Hon, 1854) Dialutacrinus tricuspidatus (de Koninck and Le Hon, 1854) Actinocrinites tripus Ehlers and Kesling, 1963 Thinocrinus tripus (Ehlers and Kesling, 1963) Actinocrinites vermiculatus Wright, 1955a Thinocrinus vermiculatus (Wright, 1955a) Actinocrinites verrucosus (Hall, 1858) Thinocrinus verrucosus (Hall, 1858) Actinocrinites zhaoae Lane et al., 1997 Incertae sedis Actinocrinites? batheri (Whidborne, 1896) Physetocrinus batheri (Whidborne, 1896) Blairocrinus grafensis Webster et al., 2004 Incertae sedis Cactocrinus excerptus (Hall, 1861) Nomen dubium Cactocrinus fosteri (McChesney, 1861) Nomen dubium Cactocrinus reticulatus (Hall, 1861) Nunnacrinus reticulatus (Hall, 1861) Cactocrinus reticulatus ovatus (Hall, 1861a) Nunnacrinus ovatus (Hall, 1861) Cusacrinus bischoffi (Miller and Gurley, 1896c) Cactocrinus bischoffi (Miller and Gurley, 1896c) Cusacrinus gracilis (Wachsmuth and Springer, 1897) Dialutocrinus gracilis (Wachsmuth and Springer, 1897) Cusacrinus hurdianus (McChesney, 1861) Cactocrinus hurdianus (McChesney, 1861) Cusacrinus imperator (Laudon, 1933) Cactocrinus imperator Laudon, 1933 Eumorphocrinus porteri (Whidborne, 1896) Incertae sedis Eumorphocrinus sp. Brower, 1970 Actinocrinites sp. (Brower, 1970) Ilmocrinus Incertae sedis Manillacrinus sp. Campbell and Bein, 1971 Nomen dubium Nunnacrinus armatus (de Koninck and Le Hon, 1854) Iotacrinus armatus (de Koninck and Le Hon, 1854) Nunnacrinus deornatus (de Koninck and Le Hon, 1854) Thinocrinus deornatus (de Koninck and Le Hon, 1854) Nunnacrinus sampsoni (Miller and Gurley, 1896c) Cusacrinus sampsoni (Miller and Gurley, 1896c) Nunnacrinus stellaris (de Koninck and Le Hon, 1854) Iotacrinus stellaris (de Koninck and Le Hon, 1854) Physetocrinus sampsoni Miller and Gurley, 1896c Thinocrinus? sampsoni (Miller and Gurley, 1896c) Physetocrinus smalleyi Weller, 1909 Blairocrinus smalleyi (Weller, 1909) 3 Rhenberg et al.—Actinocrinitid concepts

Table 1. (Continued )

Previous assignment (Webster, 2003) New assignment Sampsonocrinus cheguiaensis Webster et al., 2004 Thinocrinus cheguiaensis (Webster et al., 2004) Steganocrinus burlingtonensis Brower, 1965 Abactinocrinus burlingtonensis (Brower, 1965) Strotocrinus blairi Miller and Gurley, 1895 Teleiocrinus blairi (Miller and Gurley, 1895) Strotocrinus ornatus Miller and Gurley, 1896a Teleiocrinus ornatus (Miller and Gurley, 1896a) Strotocrinus venustus Miller and Gurley, 1893 Teleiocrinus venustus (Miller and Gurley, 1893) Teleiocrinus? sibiricus Yakovlev in Yakovlev and Ivanov, 1956 Nomen dubium three species and one genus were designated incertae sedis. This Table 2. Chronostratigraphic correlation of time units between North America is only a first step for a full understanding of the Actinocrinitidae, and Europe (from Ausich and Kammer, 2008) but it does allow for evaluation of regional and global patterns of North American generic longevity, eurytopy, and evolutionary paleoecology. Time units chronostratigraphic units European chronostratigraphic units Significant evaluation of species-level systematics remains to be 11 upper Chesterian Serpukhovian, Arnsbergian (E2) 10 middle Chesterian Serpukhovian, Pendleian (E1) completed but is far beyond the scope of this study. 9 lower Chesterian Visean, Brigantian (V3c) 8 late Meramecian Visean, Asbian (V3b) 7 early Meramecian Visean, Holkerian (V3a) Systematic paleontology 6 late Osagean Visean, Arundian (V1b, V2a) 5 late Osagean Visean, upper Chadian (V1a) 4 middle Osagean Tournaisian, lower Chadian (Tn3c) Where possible, all species were characterized using original 3 early Osagean Tournaisian, Ivorian (Tn3a-Tn3c) descriptions and illustrations, subsequent descriptions and 2 late Kinderhookian Tournaisian, late Hastarian (Tn2) illustration of type specimens, and examination of type speci- 1 early Kinderhookian Tournaisian, early Hastarian (Tn1b) mens. Complete synonymies are not provided as they are pub- lished in Webster (2003). Terminology follows Ubaghs (1978a) with modifications from Ausich et al. (1999) and open nomen- are inferred using parsimony-based character analysis of genera clature that follows Matthews (1973). using PAUP ©4.0b10 (Swofford, 2001). The actinocrinitid Carboniferous stratigraphy follows Heckel and Clayton genera are defined using discrete, non-subjective characters that (2005). Table 2 is the detailed chronostratigraphic scheme define basic and distinctive atrributes of the calyx and tegmen. showing the correlations used in this paper following Ausich Fifteen characters with 38 character states were used to analyze and Kammer (2008). the genera (Appendices 1 and 2). These characters are unordered and unweighted; search methods were heuristic with random Class Crinoidea Miller, 1821 stepwise addition. The outgroup used for this analysis are two Subclass Camerata Wachsmuth and Springer, 1885 genera from the Periechocrinidae Bronn, 1849: Periechocrinus Order Monobathrida Moore and Laudon, 1943 Morris, 1843, a Silurian and Devonian genus and Megistocrinus Suborder Compsocrinina Ubaghs, 1978b Owen and Shumard, 1852, a Devonian and Mississippian Superfamiy Periechocrinacea Bronn, 1849 genus. Characters that are used for the outgroup genera are Family Actinocrinitidae Austin and Austin, 1842 based on their type species, but the character states used for the Actinocrinitidae are based on all known species for each of Diagnosis.—The Actinocrinitidae differs from the other the genera. Thus, multiple character states are present for several three families in the Periechocrinacea (Periechocrinidae, characters in most of the genera. Amphoracrinidae, Paragaricocrinidae) in possessing a unique The subfamilies of Bowsher and Ubaghs (1978) were combination of characters that vary and in some cases overlap based on presence or absence of arm lobes, presence or absence among the four families (Ausich and Kammer, 2008, table 2). In of an anal tube, and how strongly the arms were grouped general, actinocrinitids possess a typically conical calyx that is together. None of the subfamilies form a separate clade in the higher than found in the Amphoracrinidae and Paragaricocrinidae, phylogenetic analysis (Fig. 1) completed herein. It might be but somewhat lower than those in the Periechocrinidae. The argued that the genera of the subfamily Actinocrinitinae group first primibrachial is typically hexagonal; there are few to no together, along with Cactocrinus, but the grouping is a subset of secundibrachials; there are only two plates above the hexagonal the larger clade that includes genera from the Eumorphocrininae primanal; the anal tube is rarely absent; and the number of free and Cactocrininae. As the subfamilies are scattered throughout arms ranges from 10–60, a greater range than in the other three the tree, the subfamily classification of the Treatise (Ubaghs, families. 1978b) does not recognize monophyletic clades and should be abandoned. These subfamilies were each defined on four or Remarks.—Bowsher and Ubaghs (1978) subdivided the Acti- fewer characters (Ubaghs, 1978b), in contrast to the 15 characters nocrinitidae into four subfamilies: Actinocrinitinae Austin and used in this study (Appendix 1). Austin, 1842; Eumorphocrininae Ubaghs, 1978c; Cactocrininae Wachsmuth and Springer (1897) subdivided the actinocri- Ubaghs, 1978d; and Physetocrininae Ubaghs, 1978e. A phylo- nitids into three primary groups that included: 1) genera with an genetic analysis of 20 genera from the four subfamilies (and anal tube, calyx lobed, and interradials in contact with the two outgroup genera) was performed to investigate possible tegmen; 2) genera with an anal tube, calyx not lobed, and phylogenetic relationships and to test if the previously named interradials not in contact with the tegmen; and 3) genera subfamilies represented actual clades. Phylogenetic relationships without an anal tube, regardless of other characters. Although Journal of Paleontology 89(1):1–19 4

Figure 1. Strict consensus tree based on the phylogenetic analysis of the Actinocrinitidae from PAUP© 4.0b10; 22 taxa, 15 characters, 38 character states (Appendices 1 and 2), all characters were unordered and unweighted; search methods heuristic with random stepwise addition. There were seven most parsimonious trees of length 114. Consistency index of 0.81; retention index of 0.67; rescaled consistency index of 0.54; homoplasy index of 0.75. The synapomorphies plotted at 17 nodes are as follows (see Appendix 1 for key): 1, C2, D2, E3, G0, H0, I0, K1, L0, M0, N1, O1; 2, A1, B1, F3, J0, I1, N0; 3, F2; 4, C1, J1; 5, N2; 6, A0, J2; 7, D1, E2; 8, F1; 9, C1; 10, A0, O0; 11, F2; 12, E1; 13, K0; 14, B0; 15, O2; 16, L1; 17, O1. The polytomy is where the distribution of character states for characters E, F, and L cannot be resolved by parsimony analysis. Black boxes indicate known ranges, white indicates through ranges, and grey indicates ranges that cannot be narrowed down to a time unit in the Serpukhovian. The asterisk (*) indicates an unknown range inside the Visean for Sampsonocrinus. The open circle (○) is for the subfamily Actinocrinitinae of the Treatise, the open square (□) is for the subfamily Eumorphocrininae of the Treatise, the open diamond (◊) is for the subfamily Cactocrininae of the Treatise, and the closed circle (●) is for the subfamily Physetocrininae of the Treatise. these three groups are a good general way to subdivide the family, brachials proximally and tegmen plates distally. All actinocri- they do not reflect any phylogenetic relationship between genera. nitids that have arm lobes have arms that are strongly grouped For example, Dialutocrinus lacks arm lobes but its other characters and interrays that are in contact with the tegmen. This was the group it most closely with actinocrinitids that have arm lobes, or condition that Bowsher and Ubaghs (1978) used to distinguish Eumorphocrinus with an anal tube relates more closely to two the subfamily Actinocrinitinae; the same genera are most genera that lack an anal tube, Strotocrinus and Physetocrinus,than closely related to each other as shown in the phylogenetic it does to the rest of its subfamily. tree (Fig. 1). Two other basic conditions exist among the The generic-level diagnostic characters for genera within actinocrinitids that lack arm trunks (i.e., where arms become Actinocrinitidae are defined as follows (Table 3, Appendix 1): free in a more-or-less circular pattern rather than being extended basal circlet height, radial circlet height, number of fixed into arm lobes). Most actinocrinitids that lack arm lobes have secundibrachials, highest brachitaxis (e.g., tertibrachials) in the arms that are either strongly or weakly grouped, allowing for the vertical wall of the calyx before arm lobes, number of ranges interrays to be in contact with the tegmen. In contrast, a few (rows of stacked plates) in the regular interrays, number of actinocrinitids have arm openings that are in continuous lateral ranges in posterior interray, interrays in contact with tegmen, contact, not allowing for interrays to connect with the tegmen. plating in the proximal interrays, number of plates above the Examples of these conditions are: Actinocrinites has arm lobes, primanal, arms grouped, arm lobes, presence or absence of fixed Eumorphocrinus lacks arm lobes, but has a strong grouping of interbrachials between half-rays, tegmen height compared to arm openings, Cusacrinus lacks arm lobes and only has a slight calyx, number of tegmen plates, and position of anal tube. grouping of arm openings, and Cactocrinus lacks arm lobes and One of the most distinctive characters that separate genera all the arm facets are in lateral contact with one another. within the actinocrinitids is the presence or absence of arm The actinocrinitids that have arm lobes are Aacocrinus, lobes. Arm lobes are extensions of the theca (calyx and tegmen) Abactinocrinus, Actinocrinites, Blairocrinus, Iotacrinus, away from the distal portion of the calyx, formed by fixed Sampsonocrinus, Steganocrinus, and Thinocrinus. The arm 5

Table 3. Diagnostic characters of the calyx and tegmen of genera within Actinocrinitidae

Highest brachial in vertical Number of ranges Interrays in Plating in Number of Arm Tegmen as Basal Radial Number of fixed wall of calyx before arm Number of ranges in posterior contact with proximal plates above Arms lobes Fixed intrabrachs high as Anal Tegmen Genus circlet circlet secundibrachials lobes in regular interray interray tegmen interrays primanal grouped extend between half-rays calyx opening plates Periechocrinus high low 2 Tertibrachitaxis 5+ 10+ yes 1 to 2 3 strongly no present lower eccentric many small Megistocrinus low high 2 Tertibrachitaxis 5+ 6 yes 1 to 2 3 weakly no present lower eccentric many small, few large Aacocrinus low high 1,2 Secundibrachitaxis 3 3, 4, 5 yes 1 to 2 2 strongly yes absent lower central, many eccentric medium, few

large al. et Rhenberg Abactinocrinus low high 0,1 Primibrachitaxis 2,3 4 yes 1 to 2 2 strongly yes absent lower none/ many small eccentric Actinocrinites low, high 1 Secundibrachitaxis, 3, 4, 5 4, 5 yes 1 to 3 2 strongly yes present, absent lower central, many high Tertibrachitaxis eccentric medium, many small Blairocrinus low low 1 Primibrachitaxis 2 4 yes 1 to 2 2 strongly yes absent higher central few large Cactocrinus low low 1 Secundibrachitaxis 2, 3 3 no 1 to 2 2 no no present lower, same central many medium Cusacrinus low, high 1 Secundibrachitaxis 4 6 yes 1 to 2 2 weakly no present lower, same central many high medium —

Cytidocrinus high high 0 Primibrachitaxis 4 4 yes 1 to 2 2 strongly no absent lower central many small concepts Actinocrinitid Dialutocrinus low, low, 1 Secundibrachitaxis, 3 4, 5, 6 yes 1 to 2 2 strongly, no present lower central many high high Tertibrachitaxis weakly medium Eumorphocrinus high high 2 Tertibrachitaxis 5+ 5 yes 1 to 2 2 strongly no present, absent lower central many small Glaphyrocrinus high high 2 Secundibrachitaxis 4 4, 5 yes 1 to 2 2 strongly no present lower central few large Iotacrinus low low 1, 2 Primibrachitaxis, 2, 3 3, 4 yes 1 to 2, 1 to 3 2 strongly yes present, absent lower, central, many Secundibrachitaxis higher eccentric medium Maligneocrinus low high 1 Tertibrachitaxis 5+ 5 yes 1 to 2 2 weakly no present lower ? ? Manillacrinus high high 1 Secundibrachitaxis 3 6 yes 1 to 2 2 strongly no present lower, same central many small Nunnacrinus low low, 0,1 Primibrachitaxis, 4 5 yes 1 to 2 2 strongly, no present lower, same central, many high Secundibrachitaxis weakly eccentric medium Physetocrinus low, high 1 Tertibrachitaxis, 5+ 5 yes 1 to 2 2 strongly, no present lower none/ many small high Quartabrachitaxis weakly eccentric Sampsonocrinus low, high 0, 1, 2 Primibrachitaxis, 2 3, 4 yes 1 to 2 2 strongly yes present lower central, many high Secundibrachitaxis eccentric medium, many small Steganocrinus low high 0 Primibrachitaxis 2 3 yes 1 to 2 2 strongly yes absent lower, same eccentric few large Strotocrinus high high 1 Secundibrachitaxis, 10+ ? no 1 to 2, 1 to 3 2 no no present lower none/ many small Tertibrachitaxis eccentric Teliocrinus high high 1 Tertibrachitaxis 5+ 4 yes, no 1 to 2 2 no no present lower central many small Thinocrinus low, high 1, 2 Primibrachitaxis 2,3 3, 4, 5 yes 1 to 2, 1 to 3 2 strongly yes absent lower central, many high eccentric medium, many small Journal of Paleontology 89(1):1–19 6

lobes, along with other shared characters, keep these genera are not grouped at all, and there are many more ranges in the close together in the strict consensus tree (Fig. 1, node 13). interrays than there are in Physetocrinus. Similar to Strotocrinus, Actinocrinites, Aacocrinus, Iotacrinus, and Thinocrinus Teleiocrinus has many ranges in the interrays, arms that are not are all very similar in appearance. To separate these genera the grouped, and mostly has interrays that are not in contact with the height of the radial circlet, the highest brachitaxis in the vertical tegmen (rarely the posterior interray will connect with the tegmen), wall of the calyx before the arm lobes, and the size and number but it differs by having a prominent anal tube. Eumorphocrinus is of plating on the tegmen must be examined. Actinocrinites and judged the most plesiomorphic of the actinocrinitids, having Thinocrinus both have high radial plates, but the primibrachi- various characters seen in most of the genera, but groups within taxis is the highest brachitaxis in the calyx for Thinocrinus this clade due to the synapomorphy of tertibrachitaxis as the whereas it is either the secundibrachitaxis or tertibrachitaxis for highest brachitaxis in the vertical wall of the calyx, which is found Actinocrinites. Iotacrinus has a low radial plate and Aacocrinus only in this smaller clade and the outgroup. has a few large-sized plates on the tegmen interspersed with With only seven most parsimonious trees, the phylogeny of many small-sized plates. the Actinocrinitidae is reasonably well resolved, although there Sampsonocrinus has only two ranges in the regular is a large polytomy among the genera with arm lobes. This interrays but retains many medium-sized plates on the tegmen polytomy indicates considerable homoplasy within the Actino- and fixed intrabrachials unlike the rest of the genera in the clade crinitidae. Indeed, if the genera with lobate arms are removed, of actinocrinitids with arm lobes. Blairocrinus has a tegmen that the remaining tree exhibits less homoplasy (H.I. = 0.56 [vs. is consistently higher than the calyx in all species (Iotacrinus 0.75]) and retains its previous topology. As shown by the nearly has a single species with a higher tegmen, but this is rare in the simultaneous first appearances of genera in the Tournaisian genus). Steganocrinus and Abactinocrinus only have ten free (Fig. 1), there was rapid evolutionary diversification within the arms with Steganocrinus lacking fixed secundabrachials and family, at the resolution available. This rapid diversification was Abactinocrinus having no anal tube. likely associated with predatory release following the Hangen- Other genera in the same clade containing those with arm berg Extinction event of durophagous fishes, especially the lobes are Cactocrinus, Dialutocrinus, Nunnacrinus, Cusacrinus, placoderms (Sallan et al., 2011). Evolutionary experimentation and Manillacrinus. Cactocrinus shares several characters with with calyx construction may have produced similar calyx Iotacrinus, including having low radials, but the lack of arm character states independently. Paradoxically, the oldest occur- lobes and not having interrays connected to the tegmen rences, in the Famennian, are for the highly-derived genera differentiates it. Cactocrinus groups within the lobate actino- Abactinocrinus and Physetocrinus, rather than less-derived crinitids despite lacking arm lobes because the ranges in the genera shown on the tree (Fig. 1). This suggests that the interrays are lower than those genera without arm lobes. In Actinocrinitidae may have had a more extensive evolutionary the regular interrays Cactocrinus can have either two or three history in the Devonian than currently known (Lane et al., 1997, ranges, while those without arm lobes have no fewer than 2001; Waters et al., 2003), but it only became prominent in the three. Similarly, in the posterior interray, Cactocrinus only has Tournaisian when taxa became abundant after predatory release. three ranges while the others have four or more. Dialutocrinus, The two clades recognized by cladistic analysis do not which has less than four ranges in the regular interrays, has justify naming two new subfamilies. Although mapping of either the secundibrachitaxis or tertibrachitaxis as the highest synapomorphies indicates the smaller clade is characterized by brachitaxis in the vertical wall of the calyx. Nunnacrinus has five ranges of plates in the posterior interray (node 3), and the either zero or one fixed secundibrachial and the secundibrachi- larger clade by secundibrachials as the highest brachitaxis in taxis or lower as the highest in the vertical wall of the calyx. the vertical wall of the calyx and four ranges of plates in the Nunnacrinus may have weakly grouped arms, but some species regular interrays (node 7), the morphological differences are have strongly grouped arms whereas Cusacrinus only has too subtle to recognize these clades as subfamilies within the weakly grouped arms. Cusacrinus also has more ranges in Actinocrinitidae. the interrays than Nunnacrinus. Manillacrinus only has three The Actinocrinitidae mostly existed within the Mississip- ranges in the regular interray, strongly grouped arms, and many pian, although it originated during the Famennian and ranged at small-sized plates on the tegmen. least through the Bashkirian. Geographically, the actinocrinitids Cytidocrinus and Glaphyrocrinus group together at the are recognized in North America, Western Europe, North base of the larger clade by having four ranges in all interrays; the Africa, Russia, China, Japan, and Australia. There are also only actinocrinitids with this character. Both have strongly reports of this family (Actinocrinites and Thinocrinus) from the grouped arms, but Cytidocrinus has fixed intrabrachials between Permian of Indonesia and Australia, but with such a large time half rays and many small-sized tegmen plates. Glaphyrocrinus gap between the Bashkirian and Artinskian occurrences it is lacks intrabrachials and has few large-sized tegmen plates. questionable how these few managed to survive. However, as The smaller clade of actinocrinitids includes Maligneocrinus, the characters of the Permian actinocrinitids fit well within the Physetocrinus, Strotocrinus, Teleiocrinus,andEumorphocrinus. diagnosis of the family, they are assumed to be holdovers. More Tegmen information for Maligneocrinus is not complete, but the work needs to be done on the young members of the family, but calyx shares many features with Physetocrinus. However, it is that is not within the scope of this paper. unknown whether Maligneocrinus has an anal tube or not, and the size and number of tegmen plates is not known. Genus Actinocrinites Miller, 1821 Physetocrinus and Strotocrinus both lack anal tubes. Strotocri- Type species.—Actinocrinites triacontadactylus Miller, 1821, nus does not have interrays that connect to the tegmen, the arms by subsequent designation (Wachsmuth and Springer, 1881). 7 Rhenberg et al.—Actinocrinitid concepts

Other species.—European species include: Actinocrinites ranges in posterior interray; interrays in contact with tegmen; plat- adversus (Wright, 1955a); Actinocrinites anglicus (Wright, inginproximalinterrays1to2;fixed intrabrachials between half- 1955a); Actinocrinites angustus (Wright, 1955a); Actinocrinites rays absent; arms strongly grouped; arm lobes present and extend comptus Wright, 1955a; Actinocrinites depressus Wright, laterally; tegmen lower than calyx; few large-sized and many 1955a; Actinocrinites elongatus Wright, 1955a; Actinocrinites medium-sized plates on tegmen; anal tube central or eccentric. laevissimus (Austin and Austin, 1843); Actinocrinites parkin- soni Wright, 1955a; Actinocrinites triplus Wright, 1955a; and Occurrence.—Mississippian (Tournaisian) of North America. Actinocrinites venustus (Wright, 1955a). Early Kinderhookian through middle Osagean (time units 1 to 4) of North American species include Actinocrinites grandissimus Missouri, Nevada, and New Mexico. Van Sant, 1965; Actinocrinites jugosus (Hall, 1859); Actinocrinites lobatus (Hall, 1859); Actinocrinites magnificus (Wachsmuth and Remarks.—Aacocrinus is known only from North America. Springer, 1897); Actinocrinites monticuliferus (Miller and Gurley, Brower (1967) thoroughly described this genus, which will not 1894); Actinocrinites spergenensis (Miller and Gurley, 1896a); and be repeated here. Actinocrinites sp. (Brower, 1969) [new combination]. Additionally, Actinocrinites timoricus (Wanner, 1924) is Genus Abactinocrinus Laudon and Severson, 1953 known from the Permian of Timor. Type species.—Abactinocrinus rossei Laudon and Severson, Diagnosis.—Basal circlet low or high; radial circlet high; one 1953, by original designation. fixed secundibrachial; secundibrachitaxis or tertibrachitaxis high- est brachitaxis in vertical wall of calyx; three, four, or five ranges Other species.—Abactinocrinus devonicus Waters et al., 2003 in regular interray; four or five ranges in posterior interray; inter- and Abactinocrinus burlingtonensis (Brower, 1965) [new rays in contact with tegmen; plating in proximal interrays 1 to 3; combination]. fixed intrabrachials between half-rays present or absent; arms strongly grouped; arm lobes present and extend laterally; tegmen Diagnosis.—Basal circlet low; radial circlet high; zero or one lower than or equal to calyx height; many medium-sized or many fixed secundibrachials; primibrachitaxis highest brachitaxis in small-sized plates on tegmen; anal tube central or eccentric. vertical wall of calyx; two or three ranges in regular interray; four ranges in the posterior interray; interrays in contact with Occurrence.—Mississippian (Tournaisian) through Permian tegmen; plating in proximal interrays 1 to 2; fixed intrabrachials (Artinskian) of Europe, North America, and Timor. In Europe between half-rays absent; arms strongly grouped; arm lobes Ivorian to early Chadian (time units 3 and 4) from Ireland and present and extend laterally; tegmen lower than calyx; many United Kingdom; in North America late Osagean to early small-sized plates on tegmen; anal tube absent, opening central. Meramecian (Visean) (time units 6 and 7) from Illinois, Indiana, Iowa, Kentucky, and Missouri; in Timor Artinskian. Occurrence.—Devonian (Famennian) through Mississippian (Tournaisian) of China and North America. Famennian from — Remarks. Many species previously assigned to Actinocrinites China; late Kinderhookian (time unit 2) from Montana; middle now have revised generic assignments (Table 1). Brower (1969) Osagean (time unit 4) from Iowa. tentatively assigned a specimen to the genus Eumorphocrinus, fi but the known characters of this specimen better t the genus Remarks.—Abactinocrinus is the only actinocrinitid with arm Actinocrinites. Actinocrinites sp. has a conical calyx, the half lobes that lacks an anal tube, but has previously been described rays are separated by intrabrachials, and what is known of the as having an anal tube. However, when available specimens tegmen indicates that it is a low cone. Eumorphocrinus does not were closely examined, there was no evidence of an anal tube. have these characters, having a medium bowl shaped calyx, no Abactinocrinus rossei has a small swelling at the anal opening fi xed intrabrachials between half-rays, and a low inverted bowl while A. devonicus and A. burlingtonensis have an anal opening shaped tegmen. flush with the tegmen. Genus Aacocrinus Bowsher, 1955 Genus Blairocrinus Miller, 1891 Type species.—Aacocrinus nododorsatus Bowsher, 1955, by original designation. Type species.—Blairocrinus trijugis Miller, 1891, by original designation. Other species.—North American species include Aacocrinus boonensis (Peck and Keyte, 1938); Aacocrinus chouteauensis Other species.—North American species include Blairocrinus (Miller, 1892); Aacocrinus protuberoarmatus Brower, 1967; arrosus Miller, 1892; Blairocrinus macurdai Rhenberg and Aacocrinus sampsoni (Miller and Gurley, 1895); Aacocrinus Kammer, 2013; Blairocrinus protuberus Lee et al., 2005; and senectus (Miller and Gurley, 1897); Aacocrinus spinosulus (Miller Blairocrinus smalleyi (Weller, 1909) [new combination]. and Gurley, 1893); and Aacocrinus tetradactylus Brower, 1967. Diagnosis.—Basal circlet low; radial circlet low; one fixed Diagnosis.—Basal circlet low; radial circlet high; one or two fixed secundibrachial; primibrachitaxis highest brachitaxis in vertical secundibrachials; secundibrachitaxis highest brachitaxis in vertical wall of calyx; two ranges in regular interray; four ranges in wall of calyx; three ranges in regular interray; three, four, or five posterior interray; interrays in contact with tegmen; plating in Journal of Paleontology 89(1):1–19 8

proximal interrays 1 to 2; fixed intrabrachials between half-rays tegmen; plating in proximal interrays 1 to 2; fixed intrabrachials absent; arms strongly grouped; arm lobes present and extend between half-rays present; arms not grouped; arm lobes absent; laterally; tegmen higher than calyx; few large-sized tegmen tegmen lower than or same height as calyx; many medium-sized plates; anal tube central. plates on tegmen; anal tube central.

Occurrence.—Mississippian (Tournaisian) of North America. Occurrence.—Mississippian (Tournaisian) of North America. Early Kinderhookian through middle Osagean (time units 1 to 4) of Early Kinderhookian through middle Osagean (time units 1 to 4) Kentucky, Montana, and New Mexico. from Illinois, Iowa, Missouri, New Mexico, and Utah.

Remarks.—Blairocrinus is the only actinocrinitid that has a Remarks.—Cusacrinus bischoffi, C. hurdianus,andC. imperator tegmen that is consistently higher than the calyx, which makes it are reassigned to Cactocrinus because the arms are not grouped easy to distinguish from other actinocrinitids. and the arm facets are connected to each other, not allowing the Physetocrinus smalleyi is reassigned to Blairocrinus based interrays to be in contact with the tegmen. on the characters that link it to the latter genus. These characters are the tegmen being higher than the calyx and the arms that are Genus Cusacrinus Bowsher, 1955 strongly grouped and occur in short arm lobes. In Weller’s — (1909) description, he stated that there was no anal tube, but his Type species. Actinocrinites nodobrachiatus Wachsmuth and illustrations indicate that the beginning of one located centrally Springer in Miller, 1889, by original designation. on the tegmen and broken off. — Blairocrinus arrosus is a species that is subject to much Other species. Cusacrinus arnoldi (Wachsmuth and Springer confusion, which arises from S. A. Miller assigning two in Miller, 1889); Cusacrinus asperrimus (Meek and Worthen, actinocrinitids with the species name “arrosus”: Blairocrinus 1870); Cusacrinus chloris (Hall, 1861a); Cusacrinus coelatus arrosus Miller, 1892 and Actinocrinites arrosus Miller and (Hall, 1858); Cusacrinus daphne (Hall, 1863); Cusacrinus Gurley, 1893. Without presenting a complete history of the denticulatus (Wachsmuth and Springer, 1897); Cusacrinus taxonomic confusion, Bassler and Moodey (1943) assigned the ectypus (Meek and Worthen, 1870); Cusacrinus kuenzii (Laudon 1892 species to Actinocrinites, and they left the 1893 species in et al., 1952); Cusacrinus limabrachiatus (Hall, 1861a); Cusacrinus Actinocrinites with a note stating that it probably belonged longus (Meek and Worthen, 1870); Cusacrinus ornatissmus in Cactocrinus. Webster (2003) considered both species to be in (Wachsmuth and Springer in Miller, 1889); Cusacrinus penicillus Cactocrinus (with B. arrosus also considered in the genus (Meek and Worthen, 1870); Cusacrinus sampsoni (Miller and Aacocrinus), and presumably, synonymous homonyms. How- Gurley, 1896c) [new combination]; Cusacrinus sobrinus ever, B. arrosus was correctly assigned to Blairocrinus by (Miller and Gurley, 1896c); Cusacrinus spectabilis (Miller and Miller (1892), a view endorsed by Brower (1967) and Lee et al. Gurley, 1896c) [new combination]; Cusacrinus spinotentaculus (2005). Actinocrinus arrosus Miller and Gurley, 1893 was (Hall, 1859); Cusacrinus subscitulus (Miller and Gurley, 1896c) correctly assigned to Cactocrinus. [new combination]; Cusacrinus tenuisculptus (McChesney, 1861); Cusacrinus thetis (Hall, 1861b); Cusacrinus tuberculosus Genus Cactocrinus Wachsmuth and Springer, 1897 (Wachsmuth and Springer, 1897); Cusacrinus viaticus (White, 1874). Type species.—Actinocrinus proboscidialis Hall, 1858, by ori- ginal designation. Diagnosis.—Basal circlet high or low; radial circlet high; one fixed secundibrachial; secundibrachitaxis highest brachitaxis in Other species.—Cactocrinus arrosus (Miller and Gurley, vertical wall of calyx; four ranges in regular interray; six ranges 1893); Cactocrinus bacatus Wood, 1914; Cactocrinus bischoffi in posterior interray; interrays in contact with tegmen; plating in (Miller and Gurley, 1896c) [new combination]; Cactocrinus proximal interrays 1 to 2; fixed intrabrachials between half-rays clarus (Hall, 1861a); Cactocrinus extensus Wachsmuth and present; arms weakly grouped; arm lobes absent; tegmen lower Springer, 1897; Cactocrinus fossatus (Miller, 1892); Cactocrinus than or same height as calyx; many medium-sized plates on glans (Hall, 1859); Cactocrinus hurdianus (McChesney, 1861) tegmen; anal tube central. [new combination]; Cactocrinus imperator (Laudon, 1933) [new combination]; Cactocrinus lucina (Hall, 1861b); Cacto- Occurrence.—Mississippian (Tournaisian) of North America. crinus magnidactylus Laudon and Severson, 1953; Cactocrinus Early Kinderhookian through middle Osagean (time units 1 to 4) multibrachiatus (Hall, 1858); Cactocrinus obesus (Keyes, of Iowa; Missouri; Montana; New Mexico; Nevada; Utah; and 1894); Cactocrinus opusculus (Hall, 1859); Cactocrinus platy- Alberta, Canada. brachiatus (Wood, 1914); Cactocrinus sexarmatus (Hall, 1859); Cactocrinus springeri (Rowley, 1900); and Cactocrinus Remarks.—Actinocrinites spectabilis, and A. subscitulus are thalia (Hall, 1861b). reassigned to Cusacrinus due to the lack of arm lobes, arms that are weakly grouped, appropriate number of ranges in the regular Diagnosis.—Basal circlet low; radial circlet low; one fixed interrays, and the tegmen plate sculpture. It is possible that secundibrachial; secundibrachitaxis highest brachitaxis in ver- these species are variations on one species that also includes tical wall of calyx; two or three ranges in regular interray; three C. sobrinus. They all occur in the Burlington Limestone, and the ranges in posterior interray; interrays not in contact with differences between them appear to be minor. Miller and Gurley 9 Rhenberg et al.—Actinocrinitid concepts

(1896c) even noted that they frequently found differences in the regular interrays; four, five, or six ranges in posterior interray; number of interradial plates among specimens of the same interrays in contact with tegmen; plating in proximal interrays species, which was one of the characters they used to separate C. 1to2;fixed intrabrachials between half-rays present; arms spectabilis [new comb.] and C. sobrinus. The other determining strongly or weakly grouped; arm lobes absent; tegmen lower than character between the two species is that the regular interrays are calyx; many medium-sized plates on tegmen; anal tube central. not in contact with the tegmen. Cusacrinus arnoldi and C. ornatissimus were synonymized Occurrence.—Mississippian (Tournaisian) of Europe and North with C. nodobrachiatus in Webster’s Index (2003); however, America. Ivorian and lower Chadian (time units 3–4) of Belgium, there is no explanation as to why these three species should be Ireland, England, and Wales; early Osagean (time unit 3) of Iowa. considered the same. All three species are distinguishable from each other and should remain separate. Cusacrinus nodobra- Remarks.—Cusacrinus gracilis is reassigned to Dialutocrinus chiatus has steep-sided, medium-cone shaped calyx, low basals, because the calyx shape is a bowl and not a cone. Though the and spinose plates on the tegmen, whereas C. arnoldi has a shape of the calyx is not a diagnostic character for the whole of medium-cone shaped calyx, the sides are not so steep, the basals the family, there are some genera that only have a particular are higher, and the tegmen plates are smooth. Cusacrinus shape for the calyx. Cusacrinus has a conical calyx while Dia- ornatissimus has a low-cone shaped calyx, making it different lutocrinus only has a bowl-shaped calyx. There are also only from the previous two and low basals; the tegmen is unknown. three ranges of plates in the regular interray, not four as in Nunnacrinus sampsoni has been placed in Cusacrinus Cusacrinus. The reassignment of D. gracilis extends the range because the of the high basal circlet and the highest brachitax is of Dialutocrinus to North America. in the calyx is the secundibrachitaxis. The highest brachitaxis in Actinocrinites tricuspidatus is also reassigned to Dialuto- Nunnacrinus is usuallythe primibrachitaxis and only rarely the crinus. As with Dialutocrinus gracilis, it is the medium bowl- secundibrachitaxis. Nunnacrinus has a low basal circlet. shaped calyx that makes it more closely related to Dialutocrinus Genus Cytidocrinus Kirk, 1944 because Actinocrinites is typically more conical; where it is bowl shaped, the bowl is very high. Type species.—Actinocrinus sculptus Hall, 1858, by original designation. Genus Eumorphocrinus Wright, 1955a Type species.—Eumorphocrinus erectus Wright, 1955a. Diagnosis.—Basal circlet high; radial circlet high; zero fixed secundibrachials; primibrachitaxis highest brachitaxis in ver- Other species.—Eumorphocrinus elongatus Lindley, 1979; tical wall of calyx; four ranges in regular interray; four ranges in Eumorphocrinus excelsus Wright, 1955a; and Eumorphocrinus posterior interray; interrays in contact with tegmen; plating in hibernicus Wright, 1955a. proximal interrays 1 to 2; fixed intrabrachials between half-rays absent; arms strongly grouped; arm lobes absent; tegmen lower Diagnosis.—Basal circlet high; radial circlet high; two fixed than calyx; many small-sized plates on tegmen; anal tube central. secundibrachials; tertibrachitaxis highest brachitaxis in vertical wall of calyx; five or more ranges in regular interrays; five ranges Occurrence.—Mississippian (Tournaisian) of North America. in posterior interray; interrays in contact with tegmen; plating in Early to middle Osagean (time units 3 to 4) of Iowa. proximal interrays 1 to 2; fixed intrabrachials between half-rays present or absent; arms strongly grouped; arm lobes absent; teg- — Remarks. The genus Cytidocrinus was fully described by Kirk men lower than calyx; many small-sized plates on tegmen; anal (1943, p. 263). tube central. Genus Dialutocrinus Wright, 1955a Occurrence.—Mississippian (Tournasian to Visean) of Europe Type species.—Dialutocrinus milleri Wright, 1955a. and Australia. Late Tournaisian to Visean (time units 4 to 5) of England and Ireland; Late Tournaisian (time unit 4) from New Other species.—European species include Dialutocrinus acu- South Wales. leatus (Austin and Austin, 1843); Dialutocrinus austini Ausich — and Sevastopulo, 2001; Dialutocrinus cataphractus (Austin and Remarks. The genus is described by Wright (1955a, p. 232). Austin, 1843); Dialutocrinus icosidactylus (Portlock, 1843); Genus Glaphyrocrinus Lindley, 1988 Dialutocrinus mcoyi Ausich and Sevastopulo, 2001; Dialuto- crinus polydactylus (Miller, 1821); Dialutocrinus tessellatus Type species.—Glaphyrocrinus expansus Lindley, 1988. (Phillips, 1836); and Dialutocrinus tricuspidatus (de Koninck and Le Hon, 1854) [new combination]. Other species.—Glaphyrocrinus minutus Lindley, 1988. North American species include Dialutocrinus gracilis (Wachsmuth and Springer, 1897) [new combination]. Diagnosis.—Basal circlet high; radial circlet high; two fixed secundibrachials; secundibrachitaxis highest brachitaxis in vertical Diagnosis.—Basal circlet high or low; radial circlet high or low; wall of calyx; four ranges in regular interray; four or five ranges in one fixed secundibrachial; secundibrachitaxis or tertibrachitaxis posterior interray; interrays in contact with tegmen; plating in highest brachitaxis in vertical wall of calyx; three ranges in proximal interrays 1 to 2; fixed intrabrachials between half-rays Journal of Paleontology 89(1):1–19 10

present; arms strongly grouped; arm lobes absent; tegmen lower present; arms weakly grouped; arm lobes absent; tegmen lower than calyx; few large-sized plates on tegmen; anal tube central. than calyx; tegmen plate size unknown; anal tube unknown.

Occurrence.—Mississippian (late Tournaisian, time unit 4) of Occurrence.—Mississippian (Tournaisian) of North America. Australia. Early Osagean (time unit 3) of Alberta, Canada.

Genus Iotacrinus Ausich and Sevastopulo, 2001 Remarks.—Although this genus is known only from partial crowns, the calyx shows diagnostic characteristics of actinocrinitids, con- Type species.—Actinocrinus dorsatus (de Koninck and Le Hon, firming its placement in the family. Maligneocrinus does not fitinto 1854), by original designation. other actinocrinitid genera due to its globe-shaped calyx. With little known about the tegmen, Maligneocrinus groups mostly closely Other species.—European species include: Iotacrinus armatus with Physetocrinus, although additional specimens for study may (de Koninck and Le Hon, 1894) [new combination]; Iotacrinus change the interpretation of its relationships within the family. intermedius (Wright, 1955a) [new combination]; Iotacrinus moderatus (Wright, 1955a); and Iotacrinus stellaris (de Genus Manillacrinus Campbell and Bein, 1971 Koninck and Le Hon, 1894) [new combination]. North Amer- — ican species include Iotacrinus novamexicanus Rhenberg and Type species. Cactocrinus? brownei Dun and Benson, 1920, Kammer, 2013. by original desgination. Other species.—Manillacrinus acanthus Webster and Jell, 1999. Diagnosis.—Basal circlet low; radial circlet low; one or two fixed secundibrachials; primabrachitaxis or secundabrachitaxis Diagnosis.—Basal circlet high; radial circlet high; one fixed highest brachitaxis in vertical wall of calyx; two or three ranges secundibrachial; secundibrachitaxis highest brachitaxis in ver- in regular interrays; three or four ranges in posterior interray; tical wall of calyx; three ranges in regular interray; six ranges in interrays in contact with tegmen; plating in proximal interrays posterior interray; interrays in contact with tegmen; plating in 1to2or1to3;fixed intrabrachials between half-rays present or proximal interrays 1 to 2; fixed intrabrachials between half-rays absent; arms strongly grouped; arm lobes present and extend present; arms strongly grouped; arm lobes absent; tegmen lower laterally; tegmen lower or higher than calyx; many medium- than or same height as calyx; many small-sized plates on teg- sized sized tegmen plates; anal tube central or eccentric. men; anal tube central. — Occurrence. Mississippian (Tournaisian) of Europe and North Occurrence.—Mississippian (Tournaisian) of Australia. Late America. Ivorian to early Chadian (time units 3 and 4) from Tournaisian (time units 3 or 4) of New South Wales. Belgium, Ireland, and England; early Osagean (time unit 3) from New Mexico. Remarks.—It should be noted that Bowsher (in Moore and Teichert, 1978, p. T458) erroneously referred to this genus as Remarks.—Until a recent review of the Lake Valley Formation Manillocrinus. crinoids (Rhenberg and Kammer, 2013), Iotacrinus was only known in the “Mountain Limestone” of Belgium and in the Genus Nunnacrinus Bowsher, 1955 Hook Head Formation of Ireland (Ausich and Sevastopulo, Type species.—Nunnacrinus mammillatus Bowsher, 1955, by 2001). The North American species, Iotacrinus novamexicanus, original designation. is similar to the European species but has a tegmen that is much higher and is more distinctly lobed. Other species — Nunnacrinus Actinocrinites rotundatus has been moved to Iotacrinus . North American species include dalyanus Nunnacrinus erraticus because it has a low radial circlet and two fixed secundibrachials (Miller, 1881); (Miller and Nunnacrinus foveatus in the calyx wall. Two species from Nunnacrinus have been Gurley, 1893) [new combination]; Nunnacrinus locellus moved to this genus. These are N. armatus and N. stellaris. (Miller and Gurley, 1895); (Hall, 1861b); Nunnacrinus olsoni Nunnacrinus ovatus These species have low radials which places them in Iotacrinus Ausich, 2003; (Hall, Nunnacrinus pallubrum and not Thinocrinus and fewer ranges in the interrays than 1861b) [new combination]; (Miller and Nunnacrinus pettisensis Nunnacrinus species have. Gurley, 1896b); (Miller and Gurley, 1896c); Nunnacrinus plagosus (Miller and Gurley, 1893) [new Genus Maligneocrinus Laudon, Parks, and Spreng, 1952 combination]; Nunnacrinus puteatus (Rowley and Hare, 1891); Nunnacrinus reticulatus (Hall, 1861a) [new combination]; and Type species.—Maligneocrinus medicinensis Laudon, Parks, Nunnacrinus rubra (Weller, 1909). and Spreng, 1952. One North African species is included in Nunnacrinus, N. becharensis (Webster, et al., 2004) [new combination] from Diagnosis.—Basal circlet low; radial circlet high; one fixed Algeria. secundibrachial; tertibrachitaxis highest brachitaxis in vertical wall of calyx; five or more ranges in regular interray; five ranges Diagnosis.—Basal circlet low; radial circlet low or high; zero or in posterior interray; interrays in contact with tegmen; plating in one fixed secundibrachial; primibrachitaxis or secundabrachi- proximal interrays 1 to 2; fixed intrabrachials between half-ray taxis highest brachitaxis in vertical wall of calyx; four ranges in 11 Rhenberg et al.—Actinocrinitid concepts

regular interray; five ranges in posterior interray; interrays in Nunnacrinus and Ausich and Kammer (2008) considered contact with tegmen; plating in proximal interrays 1 to 2; fixed Amphoracrinus jessieae (Miller and Gurley, 1896c) to be a intrabrachials between half-rays present; arms strongly or nomen dubium. weakly grouped; arm lobes absent; tegmen lower than or same Reassignment, herein, of Actinocrinites becharensis Web- height as calyx; many medium-sized plates on tegmen; anal tube ster et al., 2004 to Nunnacrinus extends both the geographic and central or eccentric. stratigraphic range of the latter genus. Nunnacrinus becharensis is of early Serpukovian age of western Algeria. This Algerian Occurrence.—Mississippian (Tournaisian to Serpukhovian) species differs from Actinocrinites s.s. based on the European of North America and Africa. Late Kinderhookian to early type species (Ausich and Sevastopulo, 2001; Rhenberg and Osagean (time units 2 and 3) of Iowa, Missouri, and New Kammer, 2013) because of the presence of a low basal circlet, Mexico; Serpukhovian (time unit 10) of Algeria. two plates above the first interradial in regular interrays, and a tegmen approximately as high as the calyx. Remarks.—When Bowsher (1955) named the genus Nunnacri- nus, he designated N. mammillatus as the type species. This species Genus Physetocrinus Meek and Worthen, 1869 is known only from the Nunn Member of the Lake Valley For- Type species.—Actinocrinus ventricosus Hall, 1858, by subsequent mation. However, it is a junior synonym of a previously named designation (Wachsmuth and Springer, 1881). Lake Valley Formation crinoid, N. dalyanus (Miller, 1881). Bowsher (1955, p. 20) stated that the two species are distinguish- — able from each other most readily by the number of plates that Other species. North American species include Physetocrinus separate the rays: two in N. mammillatus and one in N. dalyanus. asper (Meek and Worthen, 1870); Physetocrinus copei (Miller, This distinction is not valid as there is variation in plate number 1881); Physetocrinus dilatatus (Meek and Worthen, 1870); betweenraysinmorethan700Nunnacrinus specimens studied Physetocrinus lobatus Wachsmuth and Springer, 1897; Physeto- from the Lake Valley, including the types for N. mammillatus (see crinus majusculus Webster and Lane, 1987; and Physetocrinus Rhenberg and Kammer, 2013). Most of the specimens have both ornatus (Hall 1858). single and double plates between the rays and not just one or the Non-North American species include Physetocrinus batheri other. The other characters that Bowsher used to separate N. (Whidbourne, 1896) [new combination] and Physetocrinus mammillatus and N. dalyanus also have great variation among the brightoni Wright, 1955a. specimens studied. These include N. mammillatus being smaller in size, having a less conical calyx and tegmen, weaker plate structure Diagnosis.—Basal circlet low or high; radial circlet high; one on the calyx, and great irregularity of size of tegmen plates. All of fixed secundibrachial; tertibrachitaxis or quartibrachitaxis these features have been found to have the range of variability highest brachitaxis in vertical wall of calyx; five or more ranges judged to be within a single species of crinoid. Because of this, N. in regular interray; five ranges in posterior interray; interrays in mammillatus is a junior synonym of N. dalyanus. contact with tegmen; plating in proximal interrays 1 to 2; fixed Actinocrinus erraticus and A. plagosus are reassigned to intrabrachials between half-rays present; arms strongly to weakly Nunnacrinus because the arms are weakly grouped, do not form grouped; arm lobes absent; tegmen lower than calyx; many small- lobes, and have the appropriate number of ranges in the sized plates on tegmen; anal tube absent, opening eccentric. interrays. Although the tegmen is not known for N. plagosus, the calyx characters best fit Nunnacrinus. Occurrence.—Devonian (Famennian) through Mississippian Cactocrinus reticulatus and C. reticulatus ovatus are (Tournaisian) of North America and Europe. Famennian of Eng- reassigned to Nunnacrinus because the interrays connect with land; Ivorian (time unit 3) of Ireland; early to middle Osagean (time the tegmen, which does not occur in Cactocrinus. Furthermore, units 3–4) of Arizona, Illinois, Iowa, New Mexico, and Nevada. N. ovatus is regarded as a separate species and is not a variation of N. reticulatus, as contended by Wachsmuth and Springer Remarks.—Physetocrinus is primarily known from North (1897). Both species have a medium cone-shaped calyx, but America, although P. brightoni occurs in the Hook Head N. ovatus has a more rounded overall shapeand a nearly bowl- Formation of Ireland and P. batheri occurs in the Marwood shaped calyx. Nunnacrinus reticulatus has a medium cone- Beds of England. shaped tegmen that is about the same height as the calyx with Physetocrinus majusculus is tentatively maintained as a long spines on most of the tegmen plates, including the plates separate species from P. copei. The main difference cited by that form the base of the anal tube. Nunnacrinus ovatus has a low Webster and Lane (1987) between the two species is that bowl-shaped tegmen with short spines on a few of the tegmen P. majusculus has a more highly inflated tegmen than P. copei. plates; the spines do not appear to continue on the anal tube. A study of P. copei from the Lake Valley Formation indicates Nunnacrinus jessieae is a problematic species of Nunna- that there is a wide variation of tegmen inflation in that crinus. In 1896, Miller and Gurley described several new species (fig. 9 in Rhenberg and Kammer, 2013). Otherwise, the camerate crinoids, and two were given the trivial name differences between the two species are few. They include “jessieae”: Actinocrinites jessieae Miller and Gurley, 1896a P. majusculus having a more conical calyx, a weak overhang at and Amphoracrinus jessieae Miller and Gurley, 1896c. Webster the base of the arms, fewer plates separating the ambulacral (2003) synonymized both of these species as N. jessieae, tracts between rays and half-rays, and more uniform tegmen although they are two distinct species. Bowsher (1955) correctly plates. However, until a more detailed study of both species is assigned Actinocrinites jessieae (Miller and Gurley, 1896a) to undertaken, they are retained as separate species. Journal of Paleontology 89(1):1–19 12

With the recognition of Physetocrinus batheri (Whidborne, 1943; Steganocrinus longus Brower, 1965; Steganocrinus 1896) herein, the stratigraphic range of the genus is extended multistriatus Brower, 1965; Steganocrinus planus Brower, into the Famennian. Physetocrinus batheri is a relatively poorly 1965; Steganocrinus robustus Brower, 1965; and Steganocrinus preserved specimen. However, it belongs to the Actinocriniti- validus (Meek and Worthen, 1860). dae, and the morphology of the tegmen (Whidborne, 1898, Pl. 32, fig. 2a; Lane et al., 2001, Pl. 3, fig. 2) aligns the species with Diagnosis.—Basal circlet low; radial circlet high; zero fixed Physetocrinus. secundibrachials; primibrachitaxis highest brachitaxis in vertical wall of calyx; two ranges in regular interray; three ranges in pos- Genus Sampsonocrinus Miller and Gurley, 1895 terior interray; interrays in contact with tegmen; plating in proximal fi Type species.—Sampsonocrinus hemisphericus Miller and interrays 1 to 2; xed interbrachials between half-rays absent; arms Gurley, 1895 strongly grouped; arm lobes present and extend laterally; tegmen lower than or same height as calyx; few large-sized plates on Other species.—North American species include Sampsono- tegmen; anal tube eccentric. crinus? globosus (Wachsmuth and Springer, 1897); and Samp- sonocrinus sedaliensis (Miller and Gurley, 1895) [new Occurrence.—Mississippian (Tournaisian) of North America. combination]. Early to middle Osagean (time units 3 and 4) from Arizona, Non-North American species include Sampsonocrinus Iowa, Missouri, Nevada, and New Mexico. cannindahensis Webster and Jell, 1999 and Sampsonocrinus coplowensis (Wright, 1955a) [new combination]. Remarks.—The species Steganocrinus altus and S. longus may be junior synonyms of S. pentagonus. The descriptions by Diagnosis.—Basal circlet high or low; radial circlet high; zero, Brower (1965) indicated that the tegmen features of all three one, or two fixed secundibrachials; primibrachitaxis or secun- species are the same, and that it is only minor differences in dibrachitaxis highest brachitaxis in vertical wall of calyx; two the calyx that separates them. The differences between S. altus ranges in regular interray; three or four ranges in posterior and S. pentagonus are that the former has a more conical interray; interrays in contact with tegmen; plating in proximal dorsal cup; however, when comparing S. pentagonus (plate 92, interrays 1 to 2; fixed intrabrachials between half-rays present; figure 29) and S. altus (plate 93, figure 7), it is difficult to tell arms strongly grouped; arm lobes present and extend laterally; the two apart. Steganocrinus longus differs from S. pentagonus tegmen lower than calyx; many small-sized or many medium- by having low basals, highly tumid calyx plates, and very sized plates on tegmen; anal tube central or eccentric. high ambulacral tracts. Geographically, S. altus and S. longus occur only in the Nunn Member of the Lake Valley Formation, Occurrence.—Mississippian (Tournaisian to Visean?) of Aus- is consistent with them being morphological variants of tralia, Europe, and North America. Early Chadian (time unit 4) S. pentagonus. Because the review of the individual species is from United Kingdom; early Kinderhookian (time unit 1) from beyond the scope of this paper, they are retained as separate Missouri; Visean? from Australia. species. Steganocrinus planus Brower, 1965 is least like the other Remarks.—Actinocrinus sedaliensis and A. coplowensis are species in the genus in that all other species of Steganocrinus reassigned to the genus Sampsonocrinus because their char- have few, large plates on the tegmen, but S. planus is covered acters fit the latter genus better. Actinocrinus sedaliensis has a with numerous smaller plates. Brower (1965) noted that the low basal circlet, two ranges in the regular interrays, and an tegmen is very similar to Abactinocrinus, but all features of the eccentric anal tube and A. coplowensis has two fixed secundi- calyx compare more closely with Steganocrinus. Therefore, we brachials, two ranges of plates in the regular interrays, and follow Brower and retain S. planus within this genus. proximal plating of 1 to 2. Genus Strotocrinus Meek and Worthen, 1866b The species Sampsonocrinus? globosus is a problematic species that does not conform to any described crinoid genus. Type species.—Actinocrinites perumbrosus Hall, 1860, by Brower (1965, pg. 789) discussed this issue, suggesting it subsequent designation (Meek and Worthen, 1866a). may be a new genus of actinocrinitid. Because the species is known from a single specimen, he hesitated to name a new Diagnosis.—Basal circlet high; radial circlet high; one fixed genus and concluded that it is most similar to Sampsonocrinus secundibrachial; secundibrachitaxis or tertibrachitaxis highest and questionably placed it within this genus. With no new ’ brachitaxis in vertical wall of calyx; 10 or more ranges in regular information to add, we follow Brower s suggestion and regard interrays; number of ranges in posterior interray unknown; this species as S.? globosus. interrays not in contact with tegmen; plating in proximal inter- fi Genus Steganocrinus Meek and Worthen, 1866a rays 1 to 2 or 1 to 3; xed intrabrachials between half-rays present; arms not grouped; arm lobes absent; tegmen lower than Type species.—Actinocrinites pentagonus Hall, 1858, by original calyx; many small-sized plates on tegmen; anal tube absent, designation. opening eccentric.

Other species.—Steganocrinus altus Brower, 1965; Steganocri- Occurrence.—Mississippian (Tournaisian) of North America. nus concinnus (Shumard, 1855); Steganocrinus elongatus Kirk, Middle Osagean (time unit 4) from Iowa and Missouri. 13 Rhenberg et al.—Actinocrinitid concepts

Discussion.—Strotocrinus glyptus (Hall, 1860), is the only 1955a) [new combination]; Thinocrinus smythi Ausich and recognized species in this genus. Macurda (1974) synonymized Sevastopulo, 2001; Thinocrinus vermiculatus (Wright, 1955a) all reported members of the genus into S. glyptus, including [new combination]; and Thinocrinus wexfordensis Ausich and S. perumbrosus; stating that S. glyptus is the valid name due to Sevastopulo. page priority. Ausich and Kammer (1991b) also discussed North American species include Thinocrinus anchorensis nomenclatural history of S. glyptus; and therefore, it will not be (Webster and Lane, 1987) [new combination]; Thinocrinus repeated here. benedicti (Miller, 1892) [new combination]; Thinocrinus blairi (Miller and Gurley, 1897) [new combination]; Thinocrinus Genus Teleiocrinus Wachsmuth and Springer, 1881 botruosus (Miller and Gurley, 1895) [new combination]; Thinocrinus enigmatus (Webster and Lane, 1987) [new Type species.—Actinocrinites umbrosus Hall, 1858, by original combination]; Thinocrinus gibsoni (Miller and Gurley, 1893) designation. [new combination]; Thinocrinus lowei (Hall, 1858) [new combination]; Thinocrinus multiradiatus (Shumard, 1857) Other species.—Teleiocrinus adolescens Wachsmuth and Springer, [new combination]; Thinocrinus multiramosus altidorsatus (Row- 1897; Teleiocrinus blairi (Miller and Gurley, 1895) [new combina- ley, 1904) [new combination]; Thinocrinus pernodosus (Hall, 1858) tion]; Teleiocrinus liratus (Hall, 1859); Teleiocrinus ornatus [new combination]; Thinocrinus probolos (Ausich and Kammer, (Miller and Gurley, 1896a) [new combination]; and Teleiocrinus 1991a) [new combination]; Thinocrinus? sampsoni (Miller and venustus (Miller and Gurley, 1893) [new combination]. Gurley, 1896c) [new combination]; Thinocrinus scitulus (Miller and Gurley, 1897) [new combination]; Thinocrinus semimultiramosus Diagnosis.—Basal circlet high; radial circlet high; one fixed (Whitfield, 1900) [new combination]; Thinocrinus subpulchellus secundibrachial; tertibrachitaxis highest brachitaxis in vertical (Miller and Gurley, 1896c) [new combination]; Thinocrinus tripus wall of calyx; five or more ranges in regular interray; four ranges (Ehlers and Kesling, 1963) [new combination]; and Thinocrinus in posterior interray; interrays may or may not be in contact with verrucosus (Hall, 1858) [new combination]. tegmen; plating in proximal interrays 1 to 2; fixed intrabrachials Additionally, Thinocrinus algerianensis (Webster et al., 2004) between half-rays present; arms not grouped; arm lobes absent; [new combination]; Thinocrinus cheguiaensis (Webster et al., tegmen lower than calyx; many small-sized plates on tegmen; 2004) [new combination]; Thinocrinus combinatus (Webster, et al., anal tube central. 2004) [new combination] from Algeria; and Thinocrinus brevispina (Wanner, 1924) [new combination]; Thinocrinus brouweri (Wanner, Occurrence.—Mississippian (Tournaisian) of North America. 1924) [new combination]; Thinocrinus carinatus (Wanner, 1937) Early to middle Osagean (time units 3 and 4) of Iowa, Missouri, [new combination]; Thinocrinus dilatus (Wanner, 1924) [new Illinois, and New Mexico. combination]; Thinocrinus exornatus (Wanner, 1937) [new combination]; and Thinocrinus permicus (Wanner, 1924) Remarks.—Strotocrinus blairi and S. ornatus have been reas- [new combination] from Timor. signed to Teleiocrinus because they have an anal tube, which — is lacking in Strotocrinus. Strotocrinus venustus is Diagnosis. Basal circlet high or low; radial circlet high; one or fi reassigned to Teleiocrinus based on characters of the calyx (the two xed secundibrachials; primibrachitaxis highest brachitaxis tegmen is unknown; and therefore, it is unknown whether an in vertical wall of calyx; two or three ranges in regular interray; fi anal tube exists). The calyx of T. venustus lacks distal flaring three, four, or ve ranges in posterior interray; interrays in and has fewer ranges in the interray than Strotocrinus. Macurda contact with tegmen; plating in proximal interrays 1 to 2 or 1 to fi (1974) placed this species with Cusacrinus ectypus (Meek and 3; xed intrabrachials between half-rays present or absent; arms Worthen, 1870), but the original description (Miller and Gurley, strongly grouped; arm lobes present and extend laterally; 1893, p. 27) and illustration (pl. 6, fig. 1) indicate that this tegmen lower than or equal to calyx; many small-sized or many taxon has at least 60 arm facets that separate the calyx from medium-sized on tegmen; anal tube central. the tegmen, not allowing for the interrays to be in contact — with the tegmen. This feature distinguishes this species from Occurrence. Mississippian (Tournaisian) through Permian Cusacrinus, which has fewer arms and interrays that connect (Artinskian) of Africa, Australia, Europe, and North America. with the tegmen. Ivorian and early Chadian (time units 3 and 4) of Belgium, England, and Ireland; early Osagean to early Meramecian (time Genus Thinocrinus Ausich and Sevastopulo, 2001 unit 3 to 7) of Illinois, Indiana, Iowa, Kentucky, Missouri, New Mexico, and Nevada; Serpukhovian to middle Bashkirian of Type species.—Sampsonocrinus westheadi (Wright, 1947), by Algeria; and Artinskian of Timor. original designation. Remarks.—The separation of actinocrinitid genera on the basis Other species.—European species include Thinocrinus alatus of the strict application of diagnostic characters has resulted in (Wright, 1955a) [new combination]; Thinocrinus deornatus (de Thinocrinus being one of the most widespread actinocrinitids Koninck and Le Hon, 1854) [new combination]; Thinocrinus recognized. granulatus (Wright, 1955b) [new combination]; Thinocrinus Actinocrinites combinatus Webster et al., 2004 is reas- loricatus (Schlotheim, 1820); Thinocrinus nodosus (Wright, signed to Thinocrinus, making this the first late Mississippian 1955a) [new combination]; Thinocrinus rotundus (Wright, (Serpukhovian) species recognized in Thinocrinus. Thinocrinus Journal of Paleontology 89(1):1–19 14

combinatus has only primibrachials as part of the vertical wall 1943 Actinocrinites subpulchellus; Bassler and Moodey, of the calyx, which distinguishes it from Actinocrinites that p. 275. has either secundibrachials or higher brachitaxes fixed into the 1897 Actinocrinus griffithi Wachsmuth and Springer, calyx wall. The same criteria result in the Pennsylvanian p. 568. T. algerianensis and the six Actinocrinites species from Wanner 1897 Actinocrinus griffithi; Miller, p. 731. (1924, 1937) being reassigned to Thinocrinus, i.e. T. brevispina 1898 Actinocrinus griffithi; Weller, p. 59. (Wanner, 1924), T. broweri (Wanner, 1924), T. carinatus 1943 Actinocrinites griffithi; Bassler and Moodey, p. 270. (Wanner, 1937), T. dilatus (Wanner, 1924), T. exornatus 1973 Actinocrinites griffithi; Laudon, p. 29. (Wanner, 1937), and T. permicus (Wanner, 1924). 1977 Actinocrinites griffithi; Webster, p. 28. The holotype of Thinocrinus lowei has three plates above 1943 Actinocrinus eximius Kirk, p. 264. the primanal, which is not a characteristic of the actinocrinitids. 1973 Actinocrinites eximius; Webster, p. 35. However, Ausich and Kammer (1991a) state that all other 1897 Steganocrinus griffithi Miller and Gurley, p. 34. specimens have two plates above the primanal. Because only the 1897 Steganocrinus griffithi; Miller, p. 753. holotype has the aberrant character, T. lowei is retained in 1898 Steganocrinus griffithi; Weller, p. 600. the family. 1943 Steganocrinus griffithi; Bassler and Moodey, p. 683. The calyx characters of Thinocrinus? sampsoni [new comb.] are three ranges in the regular interray, fixed Remarks.—Kirk (1943) resassigned Actinocrinites griffithi intrabrachials between the half-rays absent, and arm lobes (Wachsmuth and Springer, 1897) to A. eximius due to name present. Physetocrinus has more ranges in the regular priority. However, this was not a necessary action as Kirk had interrays, fixed intrabrachials between half-rays, and the arms synonymized Steganocrinus griffithi with A. scitulus. In Kirk’s do not form lobes. The feature that Thinocrinus? sampsoni synonymy list for A. scitulus, he redesignated S. griffithi as shares with to Physetocrinus is the lack of an anal tube on A. griffithi, which was an unnecessary step. Due to this, A. eximius the tegmen. Only three genera of actinocrinitids are known to is considered a junior synonym to A. griffithi (Wachsmuth and lack an anal tube (Abactinocrinus, Physetocrinus, and Springer, 1897, non Miller and Gurley, 1897). Strotocrinus), but the other features of the species do not fit Despite the confusion created by Kirk, both species those genera. Thinocrinus? sampsoni is also similar to Actinocrinites griffithi and Steganocrinus griffithi are the same Abactinocrinus burlingtonensis in that both have arm lobes species which are synonymized with A. subpulchellus, now- and lack an anal tube. However, they lack other similarities. Thinocrinus subpulchellus. All three nominal species occur in The calyx shape, basal height, and interray ranges are all the same time unit (time unit 3) in the Burlington Limestone and very different and it would not make sense to assign the slight differences in size and ornamentation are not enough T.? sampsoni to the genus Abactinocrinus. Until more to justify three separate species. information can be found on this species, Thinocrinus? sampsoni will be retained with this genus. Nomina dubia.—Twenty-one species of actinocrinitids are There is some confusion with the species Actinocrinites considered to be nomina dubia because of poor illustrations or scitulus. Kirk (1943) stated that there is some question as to lack of information on the species. Most of these species are whether or not Actinocrinites sharonensis (Miller and Gurley, from the genus Actinocrinites. 1897) is a valid species but listed it as being synonymous Actinocrinites tenuistriatus (Phillips, 1841) is known only with A. scitulus. Brower (1965) stated that A. sharonensis from the stem and, therefore, cannot be placed into a genus. should be a separate species but listed A. scitulus in the Actinocrinites granulatus Goldfuss, 1831 is known from synonymy list. On the same page, Brower listed A. scitulus as a only non-descript calyx plates and at least two types of separate species. Webster (2003) followed Brower’s first columnals. Because there is such little information on the synonymy list, naming A. sharonensis as the valid name in the specimens assigned to the name, we consider it to be nomen index and A. scitulus as a junior synonym. Herein, both species dubia. As noted above, Actinocrinites granulatus Wright, are considered variations of one species. Because A. scitulus has 1955b was transferred to Thinocrinus, keeping the name valid. priority, the valid name for this species is Thinocrinus scitulus Actinocrinites cf. A. brouweri (Wanner, 1924), A. laevis [new comb.]. Miller, 1821, and Teleiocrinus? sibircus (Yakovlev in Yakovlev Webster (2003) maintained Steganocrinus griffithi Miller and Ivanov, 1956) are known only by a few plates, which and Gurley, 1897 (non Wachsmuth and Springer, 1897) as a do not give enough information to assign them confidently to separate species, even though Kirk, (1943) and Brower (1965) a genus. synonymized it with Actinocrinites scitulus. Herein, we Several species of actinocrinitids have more than just a few follow Webster (2003) in considering it as a separate species plates preserved, but lack enough information to assign them from T. scitulus; however, it is with a junior synonym of correctly anywhere, mostly due to the lack of any tegminal T. subpulchellus. information. Actinocrinites pusillus (M’Coy, 1844) has only a partial calyx that has been illustrated and described. The tegmen Thinocrinus subpulchellus (Miller and Gurley, 1896c) is missing and only one ray with partial interrays on either side are preserved.? Actinocrinites polydactylus Miller, 1821 and 1896c Actinocrinus subpulchellus Miller and Gurley, p. 13. Actinocrinites constrictus (M’Coy, 1844) are nothing but 1897 Actinocrinus subpulchellus; Miller, p. 732. internal molds of the calyx. Although individual plates can be 1898 Actinocrinus subpulchellus; Weller, p. 63. seen in the molds, there is no information on whether or 15 Rhenberg et al.—Actinocrinitid concepts

not there are arm lobes or any information on the tegmens; Cactocrinus excerptus (Hall, 1861a) has neither been therefore, there is not enough information to assign them to a genus. illustrated nor have any type specimens been located for study. Both Japanese species of Actinocrinitidae, A. higuchisawensis Until type specimens are found, C. excerptus is regarded a Minato, 1951 and A. ohmoriensis Minato, 1951, consist only of a nomen dubium. Cactocrinus fosteri (McChesney, 1861) is also calyx, which are generic enough that both could be placed into any considered a nomen dubium. This species name was assigned to number of actinocrinitid genera. Manillacrinus sp. Campbell and a specimen that did not have a tegmen and the illustration is all Bein, 1971, though in open nomenclature, only has a partial theca that remains of the original type; the types were lost in the preserved that could be part of several different genera of Chicago fire of 1871 (Wachsmuth and Springer, 1897). actinocrinitids that lack arm lobes. Actinocrinites caryocrinoides M’Coy, 1844 is based on a Incertae sedis.—Actinocrinites spinaetectus (Wanner, 1937) specimen with only four basals, making it abnormal for an from the Permian of Timor appears to be a beautiful specimen actinocrinitid. The characters that have been illustrated do not based on the illustration (Wanner, 1937, pl. 1, fig. 3–4) but agree with a single genus of actinocrinitid. The arms are broken assigning it to a genus is difficult. The calyx of A. spinaetectus off above the second primibrachial (M’Coy, 1844, pl. 26, fig. 5); fits the characters of Steganocrinus (e.g. the primibrachitaxis therefore, it is unknown if arm lobes exist or how many is highest in the vertical wall and there are no fixed secundi- secundibrachials may be fixed into the lobes and therefore the brachials) but the lack of an anal tube makes it more calyx. Unlike other actinocrinitids, there appears to be only one like Abactinocrinus. As neither Steganocrinus nor Abactino- plate in the interrays. The type is missing, and there is no crinus are known past the Tournaisian, it was decided to not information on where the specimen was collected. Therefore, force this species into a genus until the actual specimen can be further study is unlikely to occur and this species should be examined. considered nomen dubium. Eumorphocrinus porteri (Whidborne, 1896) is considered The specimen associated with Actinocrinites goldfussi incertae sedis due to poor preservation. The illustrations of (Schmidt, 1930) is a circular partial theca showing the CD Whidborne (1898, pls. 30, fig. 8; 31, figs. 1,2, and 5) and Lane interray and part of the tegmen. There is no information on the et al. (2001, pls. 1, fig. 6; 2, figs. 4 and 6; 4, fig. 1) are not clear rays, the arms or most of the tegmen, which is why this species enough to determine where this species fits within the is placed into nomen dubium. Actinocrinitidae. Lane et al. (2001) diagnosed this species as Actinocrinites marcoui Collignon, 1924 is considered a having two secundibrachials in each half-ray, which is a nomen dubium because the only illustration of it is a plate character of the genus, but this does not match the illustrations. diagram. Although the plate diagram may be accurate, it does Where the plates can be distinguished on the calyx, there is only not illustrate specimens of the species; and therefore, it cannot a single secundibrachial and not two. Therefore, more complete be compared to other species within the genus. A. decadactylus specimens are needed before this species can be assigned to (Gilbertson, 1843) is not figured, thus becoming nomen a genus. dubium. One species and one genus previously assigned to Actinocrinites skourensis (Termier, G. and Termier, H., Actinocrinitidae have characters that do not occur in the family. 1950) is illustrated only as a line drawing of a partial theca with Actinocrinites zhaoae (Lane et al., 1997) was removed from very little information on the crinoid. Unless more information Uperocrinus by Waters et al. (2003) and placed into Actinocrinites can be discovered, this species will remain a nomen dubium. based upon the hexagonal first primibrachials. However, they did Actinocrinites tenuis (de Koninck and Le Hon, 1854) is not take into account that there are three plates following the illustrated only as a part of the calyx. Based on the illustration, primanal. The Actinocrinitidae only have two plates above the A. tenuis appears to belong in the genus Aryballocrinus due to primanal. There is only one specimen of this species, thus it cannot the wide interrays and appearance of thin plates. However, be known if, like Thinocrinus lowei, three plates above the without knowing what the posterior interray looks like or primanal is an aberrant feature or the norm. Until more knowing that the plates are thin, this species is regarded as specimens can be found, this species should be excluded from nomen dubium. the Actinocrinitidae. Aacocrinus triarmatus Brower, 1967 was named for two The genus Ilmocrinus has also been determined to be incertae specimens that are poorly preserved. One specimen is very small sedis. The known specimens of I. dissymmetricus Solov’yeva, (only 4 mm), a possible juvenile, and lacks a tegmen, and the 1984 (1985) are poorly preserved, but the shape of the calyx plates other is a set of isolated arms. The preservation of these two is preserved. Actinocrinitids have a hexagonal first primibrachial specimens is too poor to have been used as the basis of a new whereas Ilmocrinus has a quadrangular one. This feature removes species and therefore A. triarmatus is designated as nomen Ilmocrinus from the Actinocrinitidae. Similarly, Blairocrinus dubium. grafensis Webster et al., 2004, from the Bashkirian of Algeria, Aacocrinus acylus Webster and Jell, 1999 is considered has a quadrangular first primibrachial. nomen dubium until the specimens can be examined. The illustrations do not show distinct plate outlines on the calyx. The illustration of the holotype (Webster and Jell, 1999, fig. 4c) Acknowledgments shows a crushed calyx with fractures that appear to follow the outline of a large calyx plate. If the facture indeed follows a For making specimens available for study, we thank K. Hollis, plate, the holotype is more likely to belong to the family Smithsonian Institution, and D.B. Blake, University of Platycrinitidae. Illinois. G.D. Sevastopulo and D.L. Meyer made important Journal of Paleontology 89(1):1–19 16

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Discussion of the classification and relations of the brachiate crinoids, and Wood, E., 1914, The use of crinoid arms in studies of phylogeny: Annals of the conclusion of the generic descriptions (1886):64–226 (separate repaged to New York. Academy of Science, v. 24, p. 1–17. continue with section 1, 139–302). Wright, J., 1947, Steganocrinus westheadi n. sp. and note on a rare crinoid and a Wachsmuth, C., and Springer, F., 1897, The North American Crinoidea blastoid from the Carboniferous Limestone of Coplow Knoll, Clitheroe: Camerata: Harvard College Museum of Comparative Zoology, Memoirs Geological Magazine, v. 84, p. 101–105. 20 and 21, 897 p. Wright, J., 1950–1960, The British Carboniferous Crinoidea. Palaeontographical Wanner, J., 1924, Die Permischen Krinoiden von Timor. 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Down, Baggy, and Pilton Beds: Proceedings of the Geological Association, v. 14, p. 371–377. H. Plating in proximal interrays: 0, 1 to 2; 1 to 3. Whidborne, G.F., 1898, A monograph of the Devonian fauna of the south of I. Number of plates above primanal: 0, 3; 1, 2. England. The fauna of the Marwood and Pilton Beds. Palaeontographical J. Arms grouped: 0, strongly; 1, no; 2 weakly. Society, Monograph 52, v. 3, p. 214–236. White, C.A., 1874, Preliminary Report upon Invertebrate Fossils Collected by K. Arm lobes present: 0, yes; 1, no. the Expeditions of 1871, 1872, and 1873. Geographical and Geological L. Fixed intrabrachials between half-rays: 0, present; 1, absent. Exploration and Surveys West of the 100th Meridian: Washington, D.C., M. Tegmen as high as calyx: 0, lower; 1, higher; 2, same. Government Printing Office, p. 1–27. Whitfield, R.P., 1900, Description of a new crinoid from Indiana: American N. Anal tube: 0, central; 1, eccentric; 2, none. Museum of Natural History, Bulletin 13, p. 23–24. O. Tegmen plates: 0, many medium; 1, many small; 2, few large. 19 Rhenberg et al.—Actinocrinitid concepts

Appendix 2. Character states for phylogenetic analysis of the Actinocrinitidae

Characters Genera A B C D E F G H I J K L M N O Periechocrinus 10 2 2 3 4 000010011 Megistocrinus 0 1 2 2 3 3 0 0 0 2 1 0 0 1 (1,2) Aacocrinus 0 1 (1,2) 1 1 (0,1,2) 0 0 1 0 0 1 0 (0,1) (0,2) Abactinocrinus 0 1 (0,1) 0 (0,1) 1 0 (0,1) 1 0 0 1021 Actinocrinites (0,1) 1 1 (1,2) (1,2,3) (1,2) 0 1 1 0 0 (0,1) 0 (0,1) (0,1) Blairocrinus 00 1 0 0 1 001001102 Cactocrinus 0 0 1 1 (0,1) 0 1 0 1 1 1 0 (0,2) 0 0 Cusacrinus (0,1) 1 1 1 2 3 0 0 1 2 1 0 (0,2) 0 0 Cytidocrinus 11 0 0 2 1 001011001 Dialutocrinus (0,1) (0,1) 1 (1,2) 1 (1,2,3) 0 0 1 (0,2) 1 0000 Eumorphocrinus 1 1 2 2 3 2 0 0 1 0 1 (0,1) 0 0 1 Glaphyrocrinus 1 1 2 1 2 (1,2) 0 0 1 0 1 0002 Iotacrinus 0 0 (1,2) (0,1) (0,1) (0,1) 0 (0,1) 1 0 0 (0,1) (0,1) (0,1) 0 Maligneocrinus 01 1 2 3 2 0012100?? Manillacrinus 1 1 1 1 1 3 0 0 1 0 1 0 (0,2) 0 1 Nunnacrinus 0 (0,1) (0,1) (0,1) 2 2 0 0 1 (0,2) 1 0 (0,2) (0,1) 0 Physetocrinus (0,1) 1 1 (2,3) 3 2 0 0 1 (0,2) 1 0021 Sampsonocrinus (0,1) 1 (0,1,2) (0,1) 0 (0,1) 0 0 1 0 0 0 0 (0,1) (0,1) Steganocrinus 0 1 0 0 0 0 0 0 1 0 0 0 (0,2) 1 2 Strotocrinus 1 1 1 (1,2) 4 ? 1 (0,1) 1 1 1 0021 Teleiocrinus 1 1 1 2 3 1 (0,1) 0 1 1 1 0001 Thinocrinus (0,1) 1 (1,2) 0 (0,1) (0,1,2) 0 (0,1) 1 0 0 1 0 (0,1) (0,1)