Middle Mississippian Disparid Crinoids from the Midcontinental United States

William I. Ausich; Thomas W. Kammer; David L. Meyer

Journal of Paleontology, Vol. 71, No. 1. (Jan., 1997), pp. 131-148.

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http://www.jstor.org Fri May 11 13:19:08 2007 J. Paleont., 71(1), 1997, pp. 131-148 Copyright @ 1997, The Paleontological Society 0022-3360/97/0071-013 1SO3.00 MIDDLE MISSISSIPPIAN DISPARID CRINOIDS FROM THE MIDCONTINENTAL UNITED STATES WILLIAM I. AUSICH, THOMAS W. KAMMER, AND DAVID L. MEYER Department of Geological Sciences, The Ohio State University, Columbus 432 10, Department of Geology and Geography, P.O. Box 6300, West Virginia University, Morgantown 26506-6300, and Department of Geology, University of Cincinnati, Cincinnati, Ohio 4522 1

ABSTRAO-Systematic restudy of Lower Mississippian disparid crinoids from the midcontinental United States results in the recognition of nine valid species and one left in open nomenclature. Although disparid crinoids are not diverse, they may be locally abundant and be the dominant crinoids in certain settings. The following species are considered valid: Catillocrinus tennesseeae Shumard, Catillocrinus wachsmuthi (Meek and Worthen), Eucatillocrinus bradleyi (Meek and Worthen), Halysiocrinus bradleyi (Meek and Worthen), Halysiocrinus cumberlandensis new species, Halysiocrinus springeri Brower, Halysiocrinus tunicatus (Hall), Synbathocrinus blairi Miller, and Synbathocrinus swallovi Hall. Several species are placed into synonymy. Catillocrinus shumardi Springer and C. turbinatus are considered junior synonyms of C. tennesseeae. Halysiocrinus granuliferus and H. perplexus are regarded as junior synonyms of H. tunicatus. Synbathocrinus angularis Miller and Gurley, S. granulatus (Wood), S. granuliferus Wetherby, S. robustus Shumard, and S. troosti (Wood) are deemed junior synonyms of S. swallovi. The youngest (Meramecian) North American calceocrinids, Halysiocrinus sp., are reported, herein, from the Harrodsburg and Salem Limestones of Indiana.

INTRODUCTION report of Halysiocrinus (or any calceocrinid) in North America ISPARID CRINOIDS are abundant, if not diverse, in several from rocks younger than the Osagean. D late Osagean and Meramecian (Mississippian) settings from the midcontinental United States, especially in mudstone STRATIGRAPHY, MATERIALS, AND METHODS and deep-water facies (Kammer and Ausich, 1987; Ausich et Late Osagean and Meramecian biostratigraphy used here fol- al., 1994). The subclass Disparida includes monocyclic crinoids lows that of Kammer et al. (1990) and Ausich et al. (1994), in with arms typically free above the aboral cup. During the Mis- which these strata were subdivided temporally into seven in- sissippian these crinoids were morphologically quite unusual tervals based on multidimensional scaling in conjunction with and interesting with the striking calceocrinids, catillocrinids, and biostratigraphic and lithostratigraphic relationships. To these synbathocrinids. The calceocrinids (Halysiocrinus)lived recum- seven intervals, an eighth is added to include the main portion bent along the sea floor with the strongly bilaterally symmetrical of the Salem Limestone in southern Indiana. Locality infor- crown folded back over the column when it was in a resting mation for these crinoids is published elsewhere: for Missouri, posture. The catillocrinids (Catillocrinus and Eucatillocrinus) Iowa, Illinois, Indiana, and northern Kentucky localities see have fine, atomous (unbranched) arms and multiple arms on Ausich and Kammer (1990, 199 la, 1991b), Ausich and Lane two radials. Synbathocrinus has five atomous arms, which are (1 980), and Kammer et al. (1 983) and for other Kentucky and quite robust. Tennessee localities see Ausich and Meyer (1988, 1992) and This study considers all late Osagean and Meramecian dis- Meyer and Ausich (1992). parids from midcontinental United States, including, among Morphometric relationships within and among species were others, those from the Mississippian stratotype area; the clastic examined using Stepwise Discriminant Function Analysis crinoidal beds in Montgomery and Monroe Counties, Indiana (SDFA) (SAS User's Guide, 1985). SDFA were run with com- and Bullitt County, Kentucky; and the Fort Payne Formation bined measurements and ratios. Measurements are listed in data of Kentucky and Tennessee. Results from this study indicate tables below and in the text; ratios are listed in the text. Results that middle Mississippian disparids have relatively long tem- are presented as plots of Canonical Variable 1 (CAN1) against poral durations and are geographically widespread. In this re- Canonical Variable 2 (CAN2) or as histograms of CAN1. gard, they are comparable to the cyathocrinoids but differ from Studied material is from the following institutions: Field Mu- advanced cladids, diplobathrids, flexibles, and monobathrids. seum of Natural History (FMNH UC), Illinois State Geological Kammer and Ausich (1993, 1996) argued that these tem~oral Survey (ISGS), Indiana University (IU), The Ohio State Uni- and geographic distributions are cdrreGted with paleoecol&cal versity (OSU), University of Illinois, Urbana (UI, UI-X), and generalizations. The disparids and cyathocrinoids are environ- U. S. National Museum of Natural History (USNM, USNM S). mental generalists; whereas the advanced cladids, diplobathrids, New material collected during this study is deposited at the U. flexibles, and monobathrids are environmental specialists S. National Museum. Classification follows Moore and Teichert (Karnmer and Ausich, 1987). (1978) with modifications of Simms and Sevastopulo (1993) A total of ten disparids are recognized from the late Osagean (note that following elevation of disparids to a subclass, orders and Meramecian of the midcontinental United States (micro- within the disparids have not been defined); terminology follows crinoids are not considered here). Nine are valid species, one is Moore (1962) and Ubaghs (1978). left in open nomenclature, and 11 species are designated herein Measurements are given in the tables for select specimens. A as junior synonyms (Table 1). One new species, Halysiocrinus complete set of measurements used in the analyses is available cumberlandensis, is named. Temporal and geographic ranges from the authors and a copy is deposited in the Springer Room, are extended for several species, for example, this is the first U. S. National Museum. 132 JOURNAL OF PALEONTOLOGY, V. 71, NO. 1, 1997

TABLE1-Summary of previously valid and new Late Osagean to early 100a-100b; VANSANT, 1964, p. 70, fig. 25.4; WEBSTER,1973, p. 145; Meramecian disparid names, as used herein. WEBSTER,1976, p. 686; UBAGHS,1978, figs. 87.2, 104.la-104.lb, 118.3; MOOREAND LANE,1978a, p. T532, fig. 326.4~;MEYER AND Svecies Status AUSICH,1983, fig. lh; WEBSTER,1986, p. 166; WEBSTER,1988, p. 92. Halysiocrinus (Calceocrinus?)perplexus SHUMARD, 1865, p. 358. Catillocrinus shumardi Springer, 1923 = C. tennesseeae Catillocrinus tennesseeae Shumard. 1865 valid Calceocrinus? perplexus (Shumard, 1865). ULRICH,1886, p. 110, 1 11, Catillocrinus turbinatus Springer, 1'923 = C. tennesseeae 113. Catillocrinus wachsmuthi (Meek and Wor- Calceocrinus perplexus (Shumard, 1865). MILLER,1889, p. 230. then, 1866) valid Halysiocrinus? perplexus (Shumard, 1865). BATHER,1893, p. 67; WEL- Eucatillocrinus bradlevi. (Meek. and Worthen. LER,1898, p. 304. 1868) valid Halysiocrinus perplexus (Shumard, 1865). SPRINGER,19 1 1, p. 194; SPR- Halysiocrinus bradleyi (Meek and Worthen, INGER,1926, p. 123, P1. 30, figs. 4-13; BASSLERAND MOODEY,1943, 1869) valid valid p. 500; UBAGHS,1953, figs. 35d-35f; MOORE,1962, p. 33, P1. 3, fig. Halysiocrinus cumberlandensis new species 6; WEBSTER,1973, p. 145; UBAGHS,1978, figs. 104.2a-104.2b, 104.3; Halysiocrinus granuliferus (Rowley, 1903) = H. tunicatus Halysiocrinus nodosus (Hall, 1860) = H. tunicatus MOOREAND LANE, 1978a, p. T532, fig. 326.4d; WEBSTER,1986, p. Halysiocrinus perplexus (Shumard, 1865) = H. tunicatus ~~~;BRowER,1990,p. 315, 317,318,figs. 7.33,7.34, 8-11, 13,Table Halysiocrinus robustus (Worthen, 1890) = H. tunicatus 1-3, 5, 6. Halysiocrinus springeri Brower, 1966 valid Calceocrinus robustus MILLER,1889, p. 230 (nomen nudum). Halysiocrinus tunicatus (Hall, 1860) valid Cheirocrinus robustus WORTHEN,1890, p. 92-93, P1. 12, fig. 7; KEYES, Halysiocrinus sp. 1894, p. 222 (as C. tunicatus). Synbathocrinus angularis Miller and Gurley, Halysiocrinus? robustus (Worthen, 1890). BATHER,1893, p. 67. 1894 = S. swallovi Synbathocrinus blairi Miller, 189 1 valid Halysiocrinus robustus (Worthen, 1890). WELLER,1898, p. 304; BASSLER Synbathocrinus granulatus (Wood, 1909) = S. swallovi AND MOODEY,1943, p. 500; MOORE,1962, p. 33; WEBSTER,1973, p. Synbathocrinus granuliferus Wetherby, 1880 = S. swallovi 145; WEBSTER,1976, p. 686; WEBSTER,1986, p. 166. Synbathocrinus robustus Shumard, 1866 = S. swallovi Halysiocrinus ventricosus (Hall, 1860). WELLER,1898 (in part), p. 304. Synbathocrinus swallovi Hall, 1858 valid Calceocrinus? granuliferus ROWLEY,1903, p. 114, P1. 35, figs. 7-9; Synbathocrinus troosti (Wood, 1909) = S. swallovi BASSLERAND MOODEY,1943, p. 346. Halysiocrinusgranulife~~~(Rowley,1903).MOORE, 1962, p. 33; WEBSTER, 1973, p. 145. Halysiocrinus springeri BROWER,1966, in part, P1. 75, fig. 23. SYSTEMATIC PALEONTOLOGY Halysiocrinus sp. MEYER,AUSICH, AND TERRY,1990, fig. 2h. Class CRINOIDEA Miller, 1821 Diagnosis. -Aboral cup adanally-abanally compressed, Subclass DISPARIDA Moore and Laudon, 1943 smooth or granulose plate sculpturing, nodes absent on A and Superfamily CALCE~CRINACEA Meek and Worthen, 1869 D radials, E superradial convex, 5-6 axil arms, no axillaries in Family CALCEOCRINIDAE Meek and Worthen, 1869 main axil series after first brachial, E arm brachials nodose, first Genus HALYSIOCRINUS Ulrich, 1886 bifurcation in E arm on brachial 7 or 8, E arm branches several Type species. -Cheirocrinus dactylus Hall, 1860; by original times exotomously, may or may not have swollen axillaries on designation. lateral arms. Description.-Crown large, recumbent on column (Figure 1.1). HALYSlOCRINUS TUNICATUS (Hall, 1860) Aboral cup large, adanally-abanally compressed. Smooth (Fig- Figures 1.1-1.5, 1.7-1.11, 1.17, 2 ure 1.2) or granulose (Figure 1.4) aboral cup plate sculpturing, Cheirocrinus tunicatus HALL, 1860, p. 124. nodes present or absent on brachials (Figure 1.7). Cheirocrinus (Calceocrinus?) tunicatus (Hall, 1860). SHUMARD,1865, p. Basal circlet two or more times as wide as high (Figure 1.3, 359. 1.10); three basal plates; elongate ligament groove along entire Calceocrinus tunicatus (Hall, 1860). HALL, 1882, fig. 1 (p. 28 1); MILLER, radial-basal articulation. 1889, p. 230; WORTHEN,1890, p. 93, P1. 12, fig. 6; KEYES, 1894, p. Radial circlet widest proximally, subtrapezoidal in shape (Fig- 222, P1. 30, fig. 4. Deltacrinus? tunicatus (Hall, 1860). UWCH, 1886, p. 110, 11 1, 113. ure 1.17); radial circlet length to proximal width ratio ranges Deltacrinus tunicatus (Hall, 1860). MILLER,1889, p. 238. from 0.58 to 1.7 1 (mean=0.99, standard deviation (SD)=0.25, Halysiocrinus tunicatus (Hall, 1860). BATHER,1893, p. 67; WELLER, number of specimens measured (n)=37); distal to proximal width 1898, p. 304; BASSLERAND MOODEY,1943, p. 501; MOORE,1962, p. ratio ranges from 0.40 to 1.07 (mean=0.55, SD=0.12, n=37). 33; VANSANT, 1964, p. 70; WEBSTER,1973, p. 145; WEBSTER,1976, A and D radials occupy majority of radial circlet, each plate p. 686; AUSICH,1978, p. 221-224; AUSICH,1980, p. 278,283; AUSICH widens distally and proximally, nodes absent, E inferradial along AND LANE,1980, p. 61; KAMMER, 1982, p. 195, 196, figs. 33x, 33y, nearly the entire radial-basal articulation (Figure 1.4, 1.9, E Table 6; KAMMER,AUSICH, AND LANE,1983, p. 55, 67; AUSICH,1983, superradial forms entire distal margin of aboral cup and bears p. 257, 276, fig. 4; KAMMER,1984, p. 117, 121, figs. 2x, 2y; KAMMER, the E-ray arm (Figure 1.2, 1.1 l), may have elongate node as on 1985a, p. 554, 555; KAMMER, 1985b, p. 41, 44; AUSICHAND LANE, E-ray brachials. B and C inferradials in sutural contact and along 1985, p. 220, 224; AUSICH,1986, p. 95; WEBSTER,1986, p. 166; BROWER,1987, p. 1003-1029, figs. 3.10, 4.10-4.12, 4.224.24, 6.15, with BC fused superradial depressed to accommodate column. Tables 2-5, 7; WEBSTER,1988, p. 92; BROWER,1990, p. 315, 317, Anal X above BC fused superradial, other details of the anal 318, figs. 7.31, 7.32, 8-11, 13, Tables, 1-3, 5, 6. series not known. Cheirocrinus nodosus HALL,1860, p. 124. Three arm-bearing rays; brachials robust, convex, E arm ex- Cheirocrinus (Calceocrinus?) nodosus (Hall, 1860). SHUMARD,1865, p. otomously branches on primibrachial 6-8, secundibrachial 3, 358. tertibrachial 3 to 4, and may branch higher; E-ray brachials Calceocrinus nodosus (Hall, 1860). HALL, 1872, P1. 6, fig. 15. commonly with elongate, horizontal nodes but nodes may be Deltacrinus nodosus (Hall, 1860). MILLER,1889, p. 238. absent (Figure 1.2). Halysiocrinus nodosus (Hall, 1860). ULRICH, 1886, p. 1 10, 1 12, 113; Lateral arms with normal bilateral heterotomy, well-devel- BATHER,1893, p. 66; SPRINGER,1926, p. 103, p. 124, 125, PI. 30, figs. 14-2 1; BAS~LERAND MOODEY,1943, p. 500; MOOREAND LAUDON, oped main axils, first brachial non-axillary then as many as 5- 1944, p. 147, PI. 55, figs. 9a-9c; UBAGHS,1953, figs. 35a-c; Moore, 6 axillaries in each arm, non-axillary plates absent in main axils 1962, p. 33, P1. 2, figs. 3a-3c, fig. 7k; YAKOVLEV,1964, p. 69, figs. (except for first brachial). The main axil may extend as high as A USICH ET AL. -MISSISSIPPIAN DISPARID CRINOIDS 133

WOURE I-Middle Mississippian halysiocrinids. 1-5. 7-11, 17. Halysiocrinus tunicatus (Hall, 1860). 1. 2. USNM S 2203, x 1.0, 1. lamview of crown with column attached, 2, dorsal view of crown showing radial circlet and Eray arm; 3.8, FMNH UC 6354 (holotype of Halysiorrinus nudosus), 3, ventral view of crown showing basal circlet and underside of arms, x 1.0,8, lateral view of partial crown, x 1.5; 4, 5, lectotype of Hafysiocrinus tunicatus, FMNH UC 12928% x 1.5, 4, dorsal view of radial circlet, 5, ventral (inner) view of radial circlet; 7, lateral view of crown, USNM S 2206 (holotype of Hafysiocrinus robustus), x 1.0; 9, lateral view of partial crown, USNM S 5084 (paratype of Halysiocrinus springml. x 1.0; 10, 11, 17, FMNH UC 12928b, c, d, pamlectotypes of Halysiocrinus tunicatus, x 1.5, 10, bad circlet, 11, 17. dorsal view of radial circlet. 6, Halysiocrinus bradleyi, holotype, ISGS 1809. x 1.5. 12-1 4, Halysiocrinus springen' Brower, 1966, USNM S 5080, holotype x 1.0.12, lateral view, 13, ventral view of crown with partial column attached, 14, dorsal view of aboral cup, slightly crushed. 15.16, Halysiocrinus sp. from the Harrodsburg Limestone, USNM S 6132, x 1.5, 15, lateral view of crown, 16, dorsal view of radial circlet. 19-22, Halysiocrinus cumberlandensis new species, x 1.5.19, dorsal view of radial circlet, USNM 483787, param, 20. dorsal view of radial circlet, USNM 483788, paratype; 21, dorsal view of radial circlet, USNM 483786, holotype; 22, dorsal view of radial circlet, USNM 483792, paratype, note the highly exaggerated nodes. zetaxil-arm and the primaxil-arm may branch as high as zeta- Columnal heteromorphic, circular, low and high columnals brachials, bihtions typically occur at intervals of three or alternate, no differentiation in proximal region except in im- four brachials. Axillaries commonly nodose; brachials approx- mature portion of column. imately as high as wide (Figure 1.8, 1.9). Discussion. -Halysiocrinus nodosus (Hall, 1860), H. robustus 134 JOURNAL OF PALEONTOLOGY, V. 71, NO. 1, 1997

(Worthen, 1890), H. perplexus (Shumard, 1865), and H. gran- plexus assignments were incorrect and 38 percent of all a priori uliferus (Rowley, 1903) are all considered junior synonyms of assignments were considered incorrect by SDFA. It should be H. tunicatus (Hall, 1860). noted that H. cumberlandensis is also not well discriminated Keyes (1894) was the first to suggest that H. tunicatus and H. from H. tunicatus on the basis of shape, however as described robustus were the same, but this revision was not followed. below, other features distinguish this new species. Results from Later, Webster (1976) suggested that both H. robustus and H. this test have F statistic probabilities that range between 0.01 nodosus were junior synonyms of H. tunicatus, and we follow and 0.04, so different populations are justifiable with these data. that here. In describing H. robustus, Worthen (1 890) illustrated A second SDFA was run on these data that only considered H. specimens of H. tunicatus and H. robustus with arms. These tunicatus and H. perplexus. In this analysis, the same two vari- specimens are similar except that H. robustus is more "robust." ables were selected, and because only two species are compared Given the range of variation now known among Keokuk Lime- only CANl is calculated. The F statistic probabilities of this stone Halysiocrinus specimens (both holotypes are from the test are 0.02 for the a priori assignments; but, again 32 percent Keokuk Limestone in the Mississippian type region), this dis- of the specimens are considered misclassified. Despite the F tinction is not sufficient to warrant a separate species. statistic for the SDFA, a separate T text on CANl values for Halysiocrinus nodosus was first described from the Edwards- H. tunicatus and H. perplexus showed that these specimens ville Formation at Crawfordsville, Indiana. However, as no belonged to the same population (F statistic probability equals Crawfordsville specimens can be confirmed as belonging to this 0.85) (Figure 2.2). species and because this species is common at Indian Creek, it The Button Mold Knob and Indian Creek populations are is assumed that the holotype probably is also from Indian Creek. separated in time, and they occurred in markedly different de- Indian Creek specimens are similar in all respects to H. robustus positional settings of the Borden deltaic complex (Ausich et al., and H. tunicatus. 1979). Some phenotypic shift is reasonable between these two Halysiocrinusperplexus was first described from Button Mold populations. Brower (1990) also considered that a plate sculp- Knob (New Providence Shale Member of the Borden Forma- turing distinction separated these species, i. e., smooth sculp- tion). Kammer (1984) designated H. perplexus as a junior syn- turing in H. tunicatus and granulose sculpturing in H. perplexus. onym of H. tunicatus, but Brower (1990, p. 3 17, 3 18) argued Fine-detail plate sculpturing in H. tunicatus is quite variable against this reassignment. Brower (1990) measured 27 speci- and is at least in part a preservational artifact. Among the H. mens ofH. perplexus from Button Mold Knob and 37 specimens tunicatus specimens examined, both smooth and granulose plate of H. tunicatus from Indian Creek. Using discriminant functions sculpturing occurs. In fact two of the syntypes of H. tunicatus for the two cases, five variables and three variables, he concluded have granulose plate sculpturing. Furthermore, specimen USNM that chi-square tests of the discriminant function scores were S 5084 from Button Mold Knob has the proximal arms pre- significantly different (Brower, 1990). Essentially, the two spe- served. Arm sculpturing and nodosity (compare Figure 1.8 to cies were regarded to be different in the relative width of the Figure 1.9) are the same as that in H. tunicatus, which further aboral cup with H. perplexus being wider. However, there was supports the similarity between Button Mold Knob and later considerable overlap in discriminant function scores between populations. In conclusion, H. tunicatus and H. perplexus are these two species, and in the three variable case only 8 1 percent two populations of one species. Although SDFA tests conclude (52 of 64) of the specimens were correctly assigned to their that two populations exist, the distinction is poorer than among respective species by discriminant function. Brower's analysis other disparid species considered below, and one third or more clearly shows that there are two different populations, but are of the a priori species assignments are considered incorrect by there two different species represented at these two locations? SDFA, again much higher than for other disparids considered Our Stepwise Discriminant Functional Analysis (SDFA) of below. When only H. tunicatus and H. perplexus are considered, Halysiocrinus (Figure 2.1) does not show significant morpho- a T Test does not distinguish two populations based on canon- logical distinction between specimens that we assign to H. tun- ical variable values (Figure 2.2). Neither the sum of all statistics icatus and H. perplexus. In the analysis, the following mea- nor inspection of specimens justifies the separation of these two surements were used as variables, along with certain ratios: species. The H. "perplexus" form simply represents a morpho- proximal width of the radial circlet (pRCW), middle width of logical extreme for this species. the radial circlet (mRCW), distal width of the radial circlet Based on Rowley's (1903) illustration and description (the (dRCW), and the radial circlet length (RCL). Ratios included type specimen is not known) of H. granuliferus, the coarse gran- height to width ratio of the radial circlet (HWRAT=RCL/ ulose plate sculpturing of this species was considered diagnostic. pRCW); the ratio of the distal to proximal radial circlet width However, as discussed above, this sculpturing, as well as other (RCRAT2=mRCW/pRCW), which measures the degree of con- characters, falls within the expected range for H. tunicatus. striction at the middle of the radial circlet. A total of 52 spec- Late Osagean and Meramecian Halysiocrinus species are dif- imens were considered in this analysis, only a representative ferentiated on the basis of aboral cup shape, aboral cup plate number are listed in Table 3. In the resultant SDFA (Figure sculpturing, number of axil arms, presence or absence of axil- 2.1), only the radial circlet median width to proximal width lanes in main axil series, nodosity of brachials, position of first ratio and the radial length to proximal width ratio were selected bifurcation in E arm, and subsequent arm branching. The char- by SDFA. Canonical Variable 1 (CAN 1) accounts for 83 percent acteristics of Halysiocrinus tunicatus are compared to other Hal- of the variance of the data and is negatively correlated with ysiocrinus species in Table 2. Specimen OSU 4769 l is an un- width ratio and positively correlated with the circlet ratio. Ca- usual specimen in that the cup is asymmetrical. It is unclear nonical Variable 2 (CAN2) accounts for 17 percent of the vari- whether this is a teratology or the result of injury. ance and is positively correlated with both variables selected. Material examined.-Syntypes of Halysiocrinus tunicatus in- Species analyzed are H. tunicatus, H. cumberlandensis n. sp., clude six specimens in lot FMNH UC 12928. These comprise H. nodosus, and H. perplexus (the latter two being junior syn- five radial circlets and one basal circlet. FMNH UC 12928a, a onyms of H. tunicatus). Although the H. perplexus points occupy radial circlet, is herein designated as lectotype (Figure 1.4, 1.5); a rather confined portion of CANl-CAN2 space, only a single the other specimens in this lot, FMNH UC 12928b-129285 are individual is clearly outside the morphospace of H. tunicatus. designated as paralectotypes (Figure 1.10, 1.1 1, 1.17). The analysis concluded that 25 percent of the a priori H. per- Specimens of the junior synonyms include the following: AUSICH ETAL.-IMISSISSIPPIAN DISPARID CRIWIDS

CAN 1 (83%) (CAN 1 81%) Halysiocriaus 7 5 4

0H. tunicaturr 0H. cumberlandensis

Catillocrinus and 0 RCRnT 2 Buca ti 11 ocrinus

,-f ,-f CAN 2 0C. tenaesseeae 0 2 3 4 (17%) C. wachsmuthi f OB.bradleyi

12 - - H. tunicatus 8 - e51 H. Tperglaxusff

CAN I Scores 2

FIGURE2-Results from Stepwise Discriminant Function Analyses (SDFA). I, SDFA for Halysiocrinus. CANl accounts for 83 percent of the variance in the data and is positively correlated with HWRAT (=RCL/pRCW, height to width ratio of aboral cup) and negatively correlated with RCRAT2 (=mRCW/pRCW, degree of constriction in middle of radial circlet); CAN2 accounts for 17 percent of the variance in the data and is positively correlated with HWRAT and RCRAT2. The filled circle is H. tunicatus holotype, half-filled circles are H. tunicatus paratypes, and circle with the vertical line is H. nodosus holotype. 2, Histogram of CANl from SDFA with only H. tunicatus and H. "perplexus"considered, CANl accounts for all of the variance. Note that the two populations form part of one normal distribution. 3, SDFA for Catilfocrinus and Eucatillocrinus. CANl accounts for 8 1 percent of the variance in the data and is positively correlated with HWRAT (=ACH/ACW2, height to width ratio of aboral cup); and CAN2 accounts for 18 percent of the variance in the data and is positively correlated with AN (arm number) and ACH (aboral cup height). The filled circle is C. tennesseeae holotype, circles with vertical lines are C. shumardi syntypes, and circle with horizontal line is a C. turbinatus syntype.

Cheirocrinus nodosus, holotype, FMNH UC 6354 (Figure 1.3), 212,887, USNM 483770-483785, USNM S 2196 (numerous previously figured specimens USNM S 2203 (Figure 1.1, 1.2); specimens), USNM S 2197 (2 specimens), USNM S 2198 (5 and Cheirocrinus robustus, holotype, USNM S 2206 (Figure specimens), USNM S 2203 (numerous specimens), USNM S 1.7). The type material for Cheirocrinus (Calceocrinus) perplex- 2204, USNM S 2205, USNM S 2207 (6 specimens), USNM S us is unknown; however, a previously figured specimen (Spring- 2208, USNM S 2209, USNM S 5084, USNM S 5968, and er, 1926, P1. 30, USNM S 2 196) was examined. Type material USNM S 5969. for Calceocrinus? granuliferus is also not known. Additionally, Occurrence. -Halysiocrinus tunicatus was originally de- one of the paratypes of Halysiocrinus springeri (USNM S 5084; scribed from the Keokuk Limestone at Warsaw and Nauvoo, Figure 1.9) is reassigned herein to H. tunicatus. Illinois. Junior synonyms were first described from material at Additional specimens assigned to this species are numerous the following localities: H. nodosus, Edwardsville Formation, and include the following: IU 15 109- 1-1 5 109-3, IU 15 109-4a- probably from Indian Creek; H. perplexus, New Providence 15109-42, IU 15109-5a-15109-5, IU 15109-5001, IU 15 109- Shale at Button Mold Knob, Kentucky; H. robustus, Keokuk 5002, IU 15109-9001-15109-9003, IU 151 13-2, IU 151 13- Limestone, Hamilton, Illinois; and I% granuliferus, New Prov- 1001-15113-1004,IU 15113-3001,IU 15113-3011,IU 15114- idence Shale at Button Mold Knob, Kentucky. 3, IU 15114-12001-151 14-12002, IU 15114-14001-15114- Halysiocrinus tunicatus is a geographically widespread spe- 14002, IU 15115-1-15115-2, IU 15121-1-15121-3, IU 15124- cies. It is now recognized from the following formations and 1, IU 15124-4-15124-5, IU 15126-201-15126-216, IU 15172- localities: 1) New Providence Shale Member of the Borden For- 5051-15 172-5052, OSU 47687-OSU 47691, USNM 37662, mation: Stones Farm, Indiana; Button Mold Knob, Kentucky USNM 37663,USNM 41,060, USNM 42,377a42,377b,USNM Solite Quarry, New Market, Kenwoad Hill, Kentucky; 2) Keo- JOURNAL OF PALEONTOLOGY, K 71, NO. 1, 1997

kuk Limestone: Keokuk, Iowa; Davids Chamber, Hamilton, Nauvoo, Niota, Illinois; Interstate 1-55 Section, Highway 61-V section, Missouri; 3) Fort Payne Formation: Cumberland Coun- ty (BlacksFerry, Highway 61-DW, Highway 6ID, Highway 6IS, and Highway 61 Ramp), Lake Cumberland (Bugwood Buildup, Cave SpringsNorth, Cave SpringsSouth, Gross Creek, Harmon Creek Buildup, Owens Branch, Pleasant Hill Buildup, Seventy- Six Falls, and Wolf CreeWCaneyFork), Kentucky;Celina Build- up, Whites Creek Springs, Tennessee; 4) Edwardsville Forma- tion: Crawfordsville, Etter Farm (probably the Edwardsville Formation), Indian Creek, Monroe Reservoir (Allens Creek Bank, Boy Scout Camp, and Waldrip Site), Stobo Bioherm, Indiana; 5) Lower Warsaw Formation Keokuk, Iowa; and 6) upper part of the Warsaw Formation: BoonvilleWest, Missouri. Measurements. -See Table 3.

HALYSI~CRINUSBRADLE~(Meek and Worthen, 1869) Figure 1.6 Calceocrinus bradleyi MEEKAND WORTHEN,1869, p. 73; MEEKAND WORTHEN,1873, p. 502-504, P1. 14, fig. 9. Deltacrinus bradleyi(Meek and Worthen, 1869). ULRICH,1886,p. 110- 113;MILLER,1889,p. 238. Halysiocrinus? bradleyi (Meek and Worthen, 1869). BATHER,1893,p. 67. ~al~siocrinusbradleyi (Meek and Worthen, 1869). WELLER,1898, p. 304;BASSLERAND MOODN,1943, p. 500;MOORE,1962, p. 33, fig. 7j; VAN SANT,1964,p. 69, 70,figs. 25.la-25.lb;WEBSTER,1973, p. 144;WOLF,1979, fig. la; WEBSTER,1986,p. 165. Diagnosis.-Aboral cup adanally-abanally compressed,coarse granulose plate sculpturing, nodes absent on A and D radials, E superradial convex, three(?) axil arms, no axillaries in main axil series after first brachial, E arm brachials not nodose, first bifurcation in E arm on brachial 5, E arm branches once iso- tomously, slightly swollen axillaries on lateral arms. Description.-Crown medium, recumbent on column. Aboral cup medium, adanally-abanallycompressed.Coarsely granulose aboral cup plate sculpturing, nodes absent on brachials (Figure 1.6). Basal circlet unknown. Radial circlet widest proximally, sub- trapezoidal in shape. Radial circlet length to proximal width $ ratio 1.06;distal to proximal width ratio 0.39. A and D radials 4 occupy majority of radial circlet, each plate widens distally and E proximally, nodes absent, E inferradial along approximately 85 90 percent of radial-basal articulation, E superradial forms entire .9 distal margin of aboral cup and bears the E arm. B and C inferradials, anal X, and anal sac unknown. -2 Three arm-bearing rays; brachials slender,convexwith aboral .-- keel, higher than wide, E arm with one isotomous branch on ? primibrachial5; Ebrachials lack nodes but aboral keel especially 2 evident on secundibrachials. E Lateral arms with normal bilateral heterotomy as known, C o well-developed main axils, first brachial non-axillary then at 9 least 3 axillaries in each arm (perhaps more), non-axillary plates 9 absent from main axils (except for first brachial). The primaxil 'E arm through tertaxil arm branch only twice to yield betabra- chials, bifurcations at intervals of three or four brachials (Figure . E -6 1.6).Axillariesnot nodose but slightly swollen; brachials higher 2 s than wide, lack nodes, with aboral keel. 2 2 Columnal longer than crown but details poorly preserved. PI Discussion. -Halysiocrinus bradleyi (Meek and Worthen, 3 1869) is a very rare but apparently a distinct species of Haly- M'8 .S 2 siocrinus. The aboral cup is indistinguishablefrom H. tunicatus, 33 because some H. tunicatus may display the coarse granulose plate sculpturing characteristic of H. bradleyi. However, the @ arms of H. bradleyi are quite distinctive, having been paedo- N - morphically simplified by both a reduction in branching and A USICH E T AL. -MISSISSIPPIAN DISPARID CRINOIDS 137 brachials that are higher than wide. Halysiocrinus bradleyi is TABLE3-Measurements for Halysiocrinus specimens (in mm). CrL, compared to other Osagean and Meramecian species in Table 2. crown length in a closed posture; pRCW, proximal radial circlet width (at articulation to basal circlet); mRCW, middle radial circlet width It is possible that the holotype of H. bradleyi has simplified (normally also the minimum width); dRCW, distal radial circlet width morphological characteristics because it is not a fully developed (E ray); RCL, radial circlet length. * Incomplete or crushed measure- adult. However, this seems very unlikely because of the size of ment; ** holotype or lectotype. This table includes only select spec- the specimen. Additional specimens of this species are necessary imens. A complete set of measurements is available from the authors, anda copy is deposited in the Springer Room, U.S. National Museum. to fully document the range of characters for H. bradleyi, and until more information is available on this species, we recom- Specimen CrL pRCW mRCW dRCW RCL mend its continued recognition as a separate species. Material examined.-The holotype and only specimen of Halysiocrinus tunicatus Halysiocrinus bradleyi, ISGS 1809. FMNH UC 12928a8* FMNH UC 12928b1 Occurrence.-Halysiocrinus bradleyi is known only from the FMNH UC 12928c1 Edwardsville Formation, Crawfordsville, Indiana. FMNH UC 12928d1 Measurements. -See Table 3. FMNH UC 12928e1 FMNH UC 63542 HALYSI~CRWUS SPRINGER1 Brower, 1966 USNM S 2204 USNMs 5084 Figure 1.12-1.14 USNM S 5968 Halysiocrinus sp. 2 SPRINGER,19 1 1, p. 194. USNM S 5969 Halysiocrinus springeri BROWER,1966, in part, p. 63 1, 632, P1. 75, figs. USNM 42377a - 1-7, (non fig. 23 = H. tunicatus);WEBSTER, 1973, p. 145. USNM 42377b - USNM 212887 50.3 Diagnosis.-Aboral cup laterally compressed and rounded in Halysiocrinus bradleyi cross section, finely granulose plate sculpturing, nodes absent ISGS 1809** 55.0 on A and D radials, E superradial convex, four axil arms, no Halysiocrinus springeri axillaries in main axil series after first brachial (except in ab- USNM S 5080** - normal specimen), E arm brachials not nodose, E arm branching USNM S 50813 - USNM S 50853 - not known, sculpturing on lateral arm not known. Description.-Crown large, recumbent on column. Aboral cup Halysiocrinus cumberlandensis USNM 483786** - large, aborally-adorally compressed or somewhat laterally com- USNM 483787 - pressed yielding more of a circular cross section for aboral cup USNM 483788 - (Figure 1.12). Smooth or finely granulose aboral cup plate sculp- USNM 483789 - turing, nodes present or absent on brachials. USNM 483790 - USNM 483791 - Basal circlet approximately twice as wide as high (Figure 1.13); USNM 483792 - three basal plates; elongate ligament groove along entire radial- USNM 483793 basal articulation; AB and CD basals form the column facet. USNM 483797 Radial circlet widest proximally, subtrapezoidal in shape; ra- USNM 483799 dial circlet length to proximal width ratio ranges from 0.85 to Halysiocrinus sp. 1.47 (mean= 1.18, SD=0.3 1, n=3); distal width to proximal width USNM S 6132** 29.8* 4.6 3.7 3.3 8.0 ratio ranges from 0.56 to 0.72 (mean=0.62; SD=0.09, n=3). A I Paralectotypes of Halysiocrinus tunicatus. and D radials occupy majority of radial circlet, extend adanally Holotype of Halysiocrinus nodosus. to produce a more circular aboral cup, each plate widens distally Paratypes of Halysiocrinus springeri. and proximally, nodes absent, E inferradial triangular along radial-basal articulation, widely separated from E superradial; designated as a paratype, USNM S 5084, by Brower (1966) is E superradial triangular, forms entire distal margin of aboral reassigned herein to H. tunicatus. cup and bears the E-ray arm, nodes absent on E superradial Occurrence.-Halysiocrinus springeri was originally described (Figure 1.14). B and C inferradials large, rectangular, in lateral from the New Providence Shale Member of the Borden For- contact, support large rectangular subanal; anal X large, rect- mation, Button Mold Knob and Bradbury Knob, Kentucky, and angular. from Stones Farm, Clark County, Indiana. No additional oc- Three arm-bearing rays; brachials robust, convex, E arm currences are known. branching unknown, proximal brachials with elongate node, Measurements. -See Table 3. wider than high. On lateral arms main axils well developed, taper adaxially, HALYSIOCRINUSCUMBERLANDENSIS new species four axillaries in each arm; the first brachial is non-axillary. Figure 1.19-1.22 Details of branching on primaxil-arm through quartaxil-arm not Diagnosis. -Aboral cup adanally-abanally compressed, known. Lateral arm brachials convex, lack nodes, wider than smooth plate sculpturing (except for nodes), large transverse high. elongate nodes on A and D radials from radial facet toward Only proximal column preserved, circular. radialhadial midline suture, E superradial very convex or with Discussion.-Halysiocrinus springeri Brower is a relatively rare transverse elongate node. Number of axil arms, main axil series, species of Halysiocrinus, but it is distinctive. It may be distin- E arm, and sculpturing on lateral arms not known. guished from H. tunicatus by having a slender aboral cup that Description.-Three arm-bearing rays; radial circlet subrect- lacks the wide base and by having only four axil arms. Halysi- angular to moderately trapezoidal (Figure 1.21); radial circlet ocrinus springeri is compared with other species in Table 2. length to proximal width ratio ranges from 0.76 to 1.08 Material examined.-Holotype of Halysiocrinus springeri (mean=0.94, SD=0.09, n= 10); distal width to proximal width USNM S 5080, and paratypes USNM S 5081 and USNM S ratio ranges from 0.43 to 0.57 (mean=0.50, SD=0.05, n=10), 5083. Other specimens, USNM S 5082 and USNM S 5085, are composed of A radial, D radial, E inferradial, and E superradial. questionably assigned to this species. One of the specimens Smooth surface sculpturing of radial plates, except for nodes. 138 JOURNAL OF PALEONTOLOGY, V. 71, NO. 1, 1997

E inferradial triangular, contains entire proximal articular fac- similar to other Salem fossils at this locality, and it is reasonable et, does not occupy entire radial circlet proximal width (Figure to conclude that specimen USNM 483809 is indeed from the 1.20). E superradial pentagonal, approximately 1.5 to 2.0 times Meramecian Salem Limestone. wider than high, occupies entire distal width of radial circlet. Occurrence.-USNM S 6 132 is from the Harrodsburg Lime- A and D radial plates of typical overall shape for Halysi- stone at Canton, Indiana; and USNM 483809 is from the Salem ocrinus. Elongate transverse node on distal portion of radial, Limestone at a roadcut at Indiana Highway 37 and Clear Creek, from radial facet to midline suture, low to protruding (Figure 0.5 mi NE of Harrodsburg, Monroe County, Indiana (SE% SEY4 1.19-1.22). Width of radial circlet less both immediately prox- sec. 20 T7N R1W Clear Creek, Indiana 7.5 min. quadrangle). imal and distal to the node. The basal circlet, arms, and column not known. Superfamily ALLAGECRINACEACarpenter and Etheridge, 188 1 Discussion.-The distinctive, protruding nodes on the radial Family CATILLQCRINIDAEWachsmuth and Springer, 1886 plates distinguish H. cumberlandensis new species from all other Genus CATILLOCRINUSShumard, 1865 Halysiocrinusspecies.Halysiocrinus curnberlandensis occurs with Type species. -Catillocrinus tennesseeae Shumard, 1865; by H. tunicatus in the Fort Payne Formation of south-central Ken- monotypy. tucky. Both species display a similar range of radial circlet shape Diagnosis. -Aboral cup medium to large size, low to flat bowl from subrectangular to trapezoidal (Figure 2. I), but the nodes or cone shape; basal plates visible or not visible in lateral view; of H. curnberlandensis easily distinguish this species. Halysi- C-ray radial with distally extending platform for articulation of ocrinus cumberlandensis is compared to other Halysiocrinus anal X; anal X triangular; anal sac medium height, wide. species in Table 2. Functional morphology. -The recumbent life style of calceo- CATILLOCRINUSTENNESSEEAE Shumard, 186 5 crinids has been discussed at length by numerous authors, most Figures 3.4-3.1 1, 3.13-3.15 recently by Brower (1966) and Ausich (1986). In a recumbent Catillocrinites tennesseeae TROOST,1849, p. 4 19 (nomen nudum); posture, the outside of the radial circlet would have lain directly TROOST,1850, p. 60 (nomen nudum). on the substratum. The elongate nodes of H. cumberlandensis Catillocrinus tennesseeae SHUMARD,1865, p. 358-359; MILLER,1889, most probably served as cleats to help anchor the crown in the p. 231; WOOD,1909, p. 24-25, P1. 9, figs. 1-3; SPRINGER,191 1, p. substratum. 194-195; SPRINGER,1923, p. 22, 26, P1. 2, figs. 1-22; BASSLERAND MOODEY,1943, p. 359; MOOREAND LAUDON, 1943, P1. 1, figs. 15a- Material examined. -Twenty-three specimens are recog- 15c; MOOREAND LAUDON,1944, p. 149, P1.55, figs. 2a-2d; UBAGHS, nized, including the holotype, USNM 483786, and 22 paratypes, 1953, figs. 44-46; WEBSTER,1973, p. 78-79; WEBSTER,1977, p. 53; USNM 483787483808. MOOREAND STRIMPLE,1978, p. T542, figs. 337.2a-c; AUSICH, 1978, Occurrence.-Halysiocrinus cumberlandensis is known from p. 225, fig. 49; AUSICHANDLANE, 1982, p. 1345, text-fig. 1; KAMMER, the following Fort Payne Formation localities: Cumberland 1982, p. 196, figs. 33z-33bb; KAMMER,1984, p. 121, figs. 2z-2bb; County (Blacks Ferry and Highway 61DW), and Lake Cum- WEBSTER,1986, p. 91; WEBSTER,1988, p. 51. berland (Bugwood Buildup, Cave Springs North, Cave Springs Catillocrinus tennesseae Shumard, 1865 [sic]. WACHSMUTHAND SPRINGER, South, Lily Creek Buildup, Owens Branch Buildup, Gross Creek, 1886,p. 268 (192), 270 (194), 272 (196); WJLWN,1948, PI. 29, figs. 6-8. Kentucky; and Celina Buildup, Tennessee. The holotype is from Catillocrinus tennesseeaea Shumard, 1865 [sic]. WELLER,1898, p. 170. Catillocrinus wachsmuthi (Meek and Worthen, 1866). AUSICH,1978, p. Cave Springs North. 227, fig. 49; AUSICH,1983, p. 276. Measurements. -See Table 3. Catillocrinus shumardi SPRINGER,1923, p. 22, 27, P1. 3, figs. 12, 13; Etymology. -The trivial name for this new species is derived BASSLERAND MOODEY,1943, p. 359; AUSICHAND LANE,1982, p. from Lake Cumberland. This crinoid is known from facies best 1345, text-fig. 1; WEBSTER,1988, p. 51. exposed along this lake. Catillocrinus turbinatus SPRINGER,1923, p. 22, 26, P1. 1, figs. 14-17; BASSLERANDMOODN, 1943, p. 359; UBAGHS,1953, fig. 43; WEBSTER, HALYSIOCRINUS Sp. 1973, p. 79; AUSICH,1978, p. 225, fig. 49; AUSICHAND LANE,1982, Figure 1.15, 1.16, 1.18 p. 1345, text-fig. 1; KAMMER,1982, p. 196; AUSICH,1983, p. 276; KAMMER,1984, p. 121; AUSICHAND LANE,1985, p. 224; WEBSTER, 1986, p. 92; WEBSTER,1988, p. 51. Discussion.-Previously, the youngest calceocrinids reported from North America were late Osagean. The next youngest re- Diagnosis.-Aboral cup large in size, low to flat bowl or cone ported occurrence is from the Permian of Russia (Moore and shape; basals not visible or only distal corners visible (in ju- Lane, 1978a). Therefore, Meramecian Halysiocrinus specimens veniles basal circlet higher); anal X much wider than C-ray arm. are worthy of note but not surprising. One specimen with arms Description.-Crown cylindrical. Aboral cup low to flat, bowl and column attached (USNM S 6 132; Figure 1.15, 1.16), housed or cone shape (Figure 3.13-3.15) (rarely concave in lateral profile in the Springer Collection and collected from the Harrodsburg (Figure 3.1 l), approximately 2.7 times higher than wide; base Limestone, Canton, Indiana, is judged to be a juvenile, due to truncate, wide; sides of aboral cup straight or convex; distal its rather small size and elongate brachials. In USNM S 6132 width:proximal width of aboral cup approximately 1.5. Aboral the radial circlet length to proximal width ratio is 8.0, and the cup plates smooth or pustulose, gently convex, sutures flush or distal to proximal radial circlet width is 0.72. Although this slightly depressed. specimen is probably a juvenile of H. tunicatus, its size, some- Basal plate circlet percentage of aboral cup height varies, in what laterally compressed calyx, elongate brachials, and reduced large specimens with convex sides typically not visible, in me- lateral arm branching make it unusual for this species. Only a dium-sized specimens with convex sides up to 10 percent of single specimen is known, so it cannot be determined with cer- aboral cup height may be the basal circlet, if sides are straight tainty whether these character differences are from being a ju- 10 percent or more may be the basal circlet, and up to 40 percent venile or a new species. Accordingly, it is treated here as Hal- may be basal circlet in juveniles; broad basal concavity. Basal ysiocrinus sp. plates may be three, unequal or partially fused (Figure 3.7,3.10). In addition, a single isolated Halysiocrinusradial plate (USNM Radial circlet height percentage typically 90 or greater. A and 483809; Figure 1.18) was collected by WIA from terra rosa on D radials large; B, C, E radials small; multiple radial facets on a bench in the Salem Limestone more than 12 m above the base A and D radials. of the Salem Limestone. The preservation of this radial plate is Articulation of anal X on a distally projecting platform on C A USZCH E T AL. -MISSISSIPPIAN DZSPARZD CRZNOZDS 139

Froma 3-Middle Mississippian catillocrinids. 1-3, Eucatillocrinus bradleyi. 1. lateral view of partial crown with anal sac broken and most arms missing, note high basal circlet, FMNH UC 5 1827A, x 1.5; 2, lateral view of aboral cup with proxistele and the proximal-most arms, FMNH UC 5 1827B, x 1.5; 3, lateral view of aboral cup with partial proxistele and the proximal-most arms, ISGS 2400, x 1.5.4-1 1.13-15, Catiffocrinus tennesseeae. 4, 5, USNM S 2224a (Catiffocrinusturbinatus syntype), x 1.5, 4, A-ray lateral view, 5 CD-intermy view, note the distal-most column is the first mesistele columnal; 6, 7, 15, USNM S 2221%holotype of Cotiffocrinustennesseeae, 6. oral view, x 1 .O, 7, basal view, x 1 .O, 15, lateral view, x 1.5; 8, lateral view of crown of juvenile, USNM S 2220 (Catiffocrinusshumardi syntype), x 1.5; ell, USNM S 2223a, x 1 .O, 9, oral view, 10, basal view, I I, lateral view, x 1.5; 13, lateral view of aboral cup, USNM S 22234 x 1.5; 14, lateral view of aboral cup, USNM S 2224b (Catillocrinus turbimtus syntype), x 1.5. 12, Catillocrinus wachsmuthi, lateral view of partial crown, USNM S 2226a, x 1.0. radial plate; anal X triangular, large, much larger than C-ray dence Shale and Fort Payne Formation), and C. shumardi (Ed- arm, higher than wide (Figure 3.5). Anal sac very wide, subcir- wardsville Formation). Springer (1923) differentiated these spe- cular in cross section, shorter than arms; plates approximately cies on the basis of aboral cup height, aboral cup width, and as high as wide. total arm number. We believe that the size difference among Arms 49 to 57 in adults (Figure 3.6, 3.9) (among known these species accounts for most of the dissimilarities (with the juveniles 24 to 44), atomous, very narrow, aborally broadly exception of C. wachsmuthi) rather than biological affinities. convex or flat. One arm on B and E radials; C radial with one Arranged from smallest to largest the species recognized by or two arms; and numerous arms on A and D radials. Brachials Springer (1923) are C. shumardi, C. wachsmuthi, C. turbinatus, approximately 3.0 times higher than wide. When in a closed and C. tennesseeae. The number and arrangements of aboral posture, arms in grooves along anal sac length. cup plates are identical in all fow species, and only C. wachs- Columnal xenomorphic. Columnals circular. Proxistele high, muthi differs in the detail of the CD interradius as discussed as many as 22 columnals (Figure 3.4); mesistele and dististele below. The temporal and geographic distributions of C. ten- not known. nesseeae and C. turbinatus have been expanded since 1923, and Discussion. -Catillocrinus and C. tennesseeae were originally they are now largely overlapping. We regard the differences recognized by Troost in his unpublished manuscript of 1849. among C. shumardi, C. turbinatus, and C. tennesseeae to rep Shumard (1 865) described these taxa giving full credit to Troost; resent intraspecific, ontogenetic variation. Accordingly, C. tur- although because Troost's names are nomina nuda, Shumard is binatus Springer, 1923 and C. shumardi Springer, 1923 are des- credited as the author of both the genus and its type species. ignated as junior synonyms of C. tennesseeae Shumard, 1865. Shumard's concept of Catillocrinus and C. tennesseeae was based Recognition that the differences among these species may on specimens collected from Button Mold Knob, Kentucky represent ontogenetic change was first proposed by Ausich and (Springer, 1923, p. 11) (Figure 3.6, 3.7). It was not until 1909 Lane (1982, p. 1345). They illustrated the distribution of aboral that this species was first illustrated by Wood, and she illustrated cup diameter, aboral cup height, and arm number among these what would have been Troost's type material (Wood, 1909, species (note that the aboral cup height range given in Springer explanation to P1.9). It is interesting to note that Troost's spec- (1923) and in Ausich and Lane (1982) of 2-22 mm for C. shu- imen, probably from Whites Creek Springs, Tennessee, is more mardi is incorrect; this is the crown height range). Aboral cup similar to the C. turbinatus concept of Springer than to the C. height and width measurements of these three species (Figure tennesseeae concept of Shumard. 4) indicate a pattern of growth with aboral cup width increasing Springer (1 923) recognized fow species of Catillocrinus from more rapidly than aboral cup height. This causes the smaller the upper part of the Burlington Limestone through late Osagean specimens to have straight-sided, cone-shaped aboral cups and strata, including C. wachsmuthi (reported from the upper Bur- older specimens to have more rounded bowl-shaped cups. Spec- lington Limestone by Springer), C. tennesseeae (New Providence imens formerly assigned to C. tennesseeae include the largest Shale and Fort Payne Formation), C. turbinatus (New Provi- specimens and are characterized by bowl-shaped aboral cups; JOURNAL OF PALEONTOLOGY, V. 71, NO. 1, 1997

muthi range is one of the juveniles from the C. shumardi type suite. The holotype of C. tennesseeae plots almost centrally in ...... the distribution, and the type specimens of C. turbinatus fall ... well within the range defined for C. tennesseeae. ti Catillocrinus tennesseeae is distinguished from the other Os- A C. vachsmvthi agean species of this genus on the basis of the basal circlet and C. tennesseeae the CD interradius. Catillocrinus tennesseeae has basals either 0 1 I I I I i not visible or only slightly visible in adults, and the anal X is 0 5 10 15 20 25 30 much wider than the C arm@); whereas C. wachsmuthi has Aboral Cup Width. basals visible in lateral view among adults, and the anal X is FIGURE4-Scatter diagram illustrating aboral cup height versus aboral the same width or narrower than the C-ray arm (Table 4). cup width for Catillocrinus tennesseeae and C. wachsmuthi;t, holo- One unusual specimen (USNM S 2223) of C. tennesseeaefrom type of C. tennesseeae, s, syntypes of C. shumardi, u, syntypes of C. Whites Creek Spring has a straight to concave (outflaring) lateral turbinatus. aboral cup profile and a reasonably narrow anal X. Although this is not typical for C. tennesseeae, it is regarded as an unusual member of this species. Only the outflared growth is problem- however, the smallest specimens in the C. tennesseeae syntype atic, because the narrowing of the anal X appears to be a result suite have a cone-shaped aboral cup more similar to Springer's of this growth. (1923) concept of C. turbinatus. Overlap of morphologies occurs Catillocrinus tennesseeae was most successful in deep-water among specimens from a single locality, which supports the settings along the eastern margin of the Eastern Interior Seaway, synonomy of these species, but it should be noted that the slight- where it was relatively common in the prodeltaic New Provi- ly smaller, more straight-sided, higher forms (the C. turbinatus dence Shale Member of the Borden Formation (Kammer, 1984; concept of Springer, 1923) occur more commonly in carbonate Kammer, 1985a) and in the fossiliferous green shale facies of facies; whereas the large extremely low, convex-sided forms the Fort Payne Formation (Ausich and Meyer, 1990). In other (closer to the holotype of C. tennesseeae) are more typical in facies it is relatively rare with only juveniles typically known. shale facies. All of the material of C. shumardi is from the Material examined. -The holotype of Catillocrinus tennes- Edwardsville Formation at Indian Creek, Indiana; and these are seeaeis USNM S 222 la (Figure 3.6,3.7). Type material ofjunior all very small juveniles, which accounts for their somewhat synonyms are as follows: Catillocrinus turbinatus, syntypes distinctive morphology. USNM S 2224 (2 specimens) (Figure 3.4, 3.5, 3.14); Catillo- SDFA of all three catillocrinid species displays good mor- crinus shumardi, syntypes USNM S 2220 (5 specimens) (Figure phometric separation (Figure 2.3). A combination of measure- 3.8). Additional material includes the following: IU 8266-18- ments and measurement ratios were used. Measurements are 8266-22, IU 15172-5056, IU 15171-5057,OSU 47697, USNM aboral cup height (ACH), aboral cup width along A-CD axis 3991 1 (4 specimens), USNM 4838 10-4838 16, USNM S 2221 (ACWl), aboral cup width perpendicular to ACWl (ACW2), (14 additional specimens), USNM S 2222, USNM S 2223a, and basal circlet height (BCH), and arm number (AN). Ratios in- USNM S 2223b. clude the height to width ratio of the aboral cup (HWRAT=ACW Occurrence.-Shumard's description of Catillocrinus tennes- ACW2); relative contribution of the basal circlet to the aboral seeae was based on material from the New Providence Shale, cup height (BCPER=BCH/ACH); and ratio between aboral cup Button Mold Knob, Kentucky. Junior synonyms were originally widths (ACRAT=ACW2/ACW I), which is the measure of the described from the following: C. turbinatus, New Providence deviation of the cup outline from a circle. A total of 29 speci- Shale, Button Mold Knob, Kentucky, and Fort Payne Forma- mens were considered in this analysis, and only a representative tion, Whites Creek Springs, Tennessee; C. shumardi, Edwards- number of these are listed in Table 5. This analysis identified ville Formation, Indian Creek, Indiana. This species now is the following characters as significant; cup height:width ratio, recognized from the following formations and areas: 1) New arm number, A-CD cup width: BC-DE cup width, and cup Providence Shale Member of the Borden Formation: Button height. CAN1 accounts for 81 percent of the variance and is Mold Knob, Kentucky Solite Quarry, Kentucky; 2) Edwardsville controlled primarily by the cup height:width ratio, and CAN2 Formation: Indian Creek, Monroe Reservoir (Allens Creek and accounts for 19 percent of the variance and is largely a function Causeway Section), and Etter Farm (probably the Edwardsville of the remaining three variables. In this analysis only one of 29 Formation), Indiana; and 3) Fort Payne Formation: Cumber- specimens was considered misclassified. Results from this test land County (BlacksFerry, Highway 6 1DW, and Highway 61D), have F statistic probabilities of 0.0001. Eucatillocrinus bradleyi Lake Cumberland (Bugwood Buildup, Cave Springs North, Cave is clearly separated from the two species of Catillocrinus, and Springs South, Greasy Creek Buildup, Harmon Creek, Owens the specimen of C. tennesseeae that overlaps with the C. wachs- Branch, Pleasant Hill Buildup), Kentucky, and Celina Buildup,

TABLE4-Matrix of adult charactersfor Late Osageancatillocrinidspecies. As discussed in text,juveniles of C. tennesseeaemay be morphologically similar to other species.

- C radial Anal sac extension Anal x Anal x wider high and Species Aboral cup size Aboral cup shape Basals visible for anal x triangular than C am narrow C. tennesseeae Large Low bowl* No or distal Yes Yes Yes No corners only* C. wachsmuthi Medium Low bowl or Yes Yes Yes No No cone E. bradleyi Small Medium to Yes No No Yes Yes high cone * In juveniles the aboral cup is conical and basals are clearly visible. AUSICH E T AL. -MISSISSIPPIAN DISPARID CRINOIDS 141

5 miles west of Edgemead Station, and Whites Creek Spring, TABLE5-Measurements of Catillocrinus and Eucatillocrinus speci- Tennessee. mens. CrH, crown height; ACH, aboral cup height; ACW1, A-CD Measurements.-See Table 5. aboral cup width; ACW2, aboral cup width perpendicular to ACW 1; BCH, basal circlet height; AN, arm number. All measurements in mm, * denotes measurement incomplete or specimen crushed, ** CATILLOCRINUSWACHSMUTHI (Meek and Worthen, 1866) denotes primary type. This table includes only select specimens. A Figure 3.1 2 complete set of measurements is available from the authors, and a Synbathocrinus (Nernatocrinus) wachsmuthi MFEKAND WORTHEN,1866, copy is deposited in the Springer Room, U.S. National Museum. p. 25 1-252. Catillocrinus wachsrnuthi (Meek and Worthen, 1866). MEEKAND WOR- Species/Specimen CrH ACH ACWl ACW2 BCH AN THEN,1868a, p. 465-466, P1. 18, fig. 5; WACHSMUTHAND SPRINGER,Catillocrinus tennesseeae 1885, P1. 5, figs. 15, 15a, 16; WACHSMUTHAND SPRINGER,1886, p. USNMS2221a** - 9.0 20.8 22.8 0.1 49 269 (193)-272 (196); MILLER,1889, p. 231, fig. 263; WACHSMUTH USNM S 22201 21.9 2.0* 3.3* 5.5* 0.8* 29 AND SPRINGER,1897, P1. 8, fig. 14; WELLER,1898, p. 170; SPRINGER, USNM S 22201 25.1 2.6* 4.3* 5.6* 1.0* 23 1923, p. 22, 26-27, P1. 3, figs. 1-1 1; BASLER AND MOODEY,1943, p. USNM S 22201 16.3* 1.9* 2.0* 4.4* 0.7* 24 359; MOOREAND LAUDON,1944, p. 149, P1. 55, fig. 1; VAN SANT, USNM S 222 1 - 9.1 23.3 26.0 0.0 57 1964, p. 68, figs. 24.2a-24.2b; WEBSTER,1973, p. 79; MOOREAND USNM S 2221 - 7.5 20.2 22.4 0.0 51 USNM S 2221 7.1 19.8 21.1 0.0 53 STRIMPLE,1978, p. T542, figs. 337.2d-2e; AUSICHAND LANE,1982, - USNM S 2221 - 7.3 21.8 25.2 0.0 55 p. 1345, text-fig. 1; WEBSTER,1986, p. 92; WEBSTER,1988, p. 51. USNM S 2221 - 9.1 24.8 26.0 0.9 56 Diagnosis.-Aboral cup medium in size, low bowl or cone USNM S 22242 24.5* 5.6* 6.4* 11.0* 1.2* 41 USNM S 22242 - 4.9 10.0 11.2 1.2 - shape; basals visible; anal X as wide as or narrower than C-ray arm. Catillocrinus wachsmuthi Discussion.-Catillocrinus wachsmuthi is a reasonably rare USNM S 2226 43.5* 4.3 - 12.8 1.0 - USNM S 2226 - 3.0 6.2 6.3 0.9 26 Burlington crinoid, only known from a handful of specimens. USNM S 2226 29.4* 4.7 11.3 12.4 1.3 39 It is distinguished from C. tennesseeae on the basis of the CD USNM S 2226 20.7* 2.1 5.3 5.7 0.7 25 interradius and the basal circlet, as discussed above and in Table Eucatillocrinus bradleyi 4. It is assumed that the available C. wachsmuthi specimens are ISGS 2400** - 3.9 - - 1.0 38* adults. Based solely on aboral cup height and width (Figure 4), FMNH UC 51827a - 6.0 5.0 - 1.7 - C. wachsmuthifalls within the range of C. tennesseeae. However, FMNHUC 51827b - 5.2 4.7 6.5* 1.8 39* SDFA of C. wachsmuthi indicates that this species has a dis- USNM S 2215 29.8* 4.8 - - 1.3 - USNM S 2215 - 5.6 5.4* 6.4* 2.1 43 tinctive morphology. The only specimen of C. tennesseeae that USNM S 22 15 - 5.4 5.5* 8.4* 1.6 46 plots among the C. wachsmuthi specimens is a very small spec- USNM S 2215 77.6* 6.6 - - 2.0 - imen that plots nearest the smallest C. wachsmuthi. All other USNM S 2215 65.2* 6.6 - - 1.9 - C. tennesseeae, most of which are larger, plot more positively I Catillocrinus shumardi syntype. on CAN2. Alternatively, larger C. wachsmuthi plot more neg- Catillocrinus rurbinarus syntype. atively on CAN2. This morphometric distinction as well as the differences in the posterior interray are judged sufficient to retain C. wachsmuthi as a valid species at this time. Eucatillocrinus bradleyi (Meek and Worthen, 1868). SPRINGER,1923, Material examined. -The Meek and Worthen (1866) type p. 19, 23, PI. 3, figs. 14-17; BASSLERAND MOODEY,1943, p. 468; material for Catillocrinus wachsmuthi is unknown. Six speci- MOOREAND LAUDON,1944, p. 149, P1. 55, fig. 3; VAN SANT, 1964, mens are in the lot of material studied by Springer (1 923), USNM p. 68-69, P1. 1, figs. 1-2, figs. 24.la-24. lb; WEBSTER,1973, p. 127; S 2226. STRIMPLEET AL., 197 1, p. 13; WEBSTER,1977, p. 82-83; MOOREAND Occurrence.-Catillocrinus wachsmuthi is known from the STRIMPLE,1978, p. T545, fig. 340.2; WOLF,1979, fig. 1 (unnumbered); upper part of the Burlington Limestone at Burlington, Iowa WEBSTER,1986, p. 144. (middle Osagean). Diagnosis.-See generic diagnosis. Measurements.-See Table 5. Description.-Crown narrow, cylindrical. Aboral cup medi- um to high cone shape (Figure 3.2), approximately 1.3 times Genus EUCA~LLOCRINUSSpringer, 1923 higher than wide; base truncate, wide; sides of aboral cup straight, Type species. -Catillocrinus bradleyi Meek and Worthen, widen distally; distal width:proximal width of aboral cup ap- 1868, by original designation. proximately 1.4. Aboral cup plates smooth, gently convex, su- Diagnosis. -Aboral cup medium size, medium to high cone tures flush. shape; basal plates visible in lateral view; C radial lacking dis- Basal plate circlet 27 to 38 percent of aboral cup height tally extending platform for articulation of anal X; anal X rect- (mean-3 1, n=7); expands in width distally; no basal concavity. angular; anal sac very high, slender. Basal plates may be three, unequal, or partially fused. Radial Discussion.-Although a monospecific genus, Eucatillocrinus circlet height approximately 69 percent of cup height; expands displays consistent, distinctive differences in the construction distally at same rate as distal expansion of basal circlet. A and of the aboral cup, arms, and anal sac that warrant its separation D radials large; B, C, E radials small; multiple radial facets on from Catillocrinus as a separate genus (Table 4). This crinoid A and D radials. is reasonably rare in the Crawfordsville and Indian Creek faunas. Articulation of anal X level with arm articulations; anal X rectangular, large, much larger than C-ray arm, as wide as high EUCA~LLOCRINU~BRADLEYI (Meek and Worthen, 1868) or slightly wider than high (Figure 3.3). Anal sac slender, sub- Figure 3.1-3.3 circular in cross section, much longer than arms; plates higher than wide (Figure 3.1). Catillocrinus bradleyi MEEKAND WORTHEN,1868b, p. 343; MEEKAND WORTHEN,1873, p. 504-505, P1. 14., figs. 10a, lob; WACHSMUTHAND Arms 39 to 46, atomous, very narrow, aborally broadly con- SPRINGER,1886, p. 269 (193), 272 (196); MILLER,1889, p. 231; WEL- vex or flat. One arm on B, C, and E radials; numerous arms on LER, 1898, p. 170; L NOT, 1948, p. 62, fig. 80; YAKOVLEV,1964, p. A and D radials. First primibrachial approximately 3.0 times 64, fig. 84. higher than wide; second primibrachial approximately 1.5 times 142 JOURNAL OF PALEONTOLOGY, V. 71, NO. 1, 1997

wider than high. When in a closed posture, arms may fit into Diagnosis. -Large aboral cup, medium cone-shaped aboral short grooves along the proximal portion of the anal sac. cup with sides either straight or concave in lateral view; low Columnal xenomorphic. Columnals circular, lumen penta- basal circlet; gently to strongly convex radial plates, radia1:radial lobate. Proxistele high, composed of as many as 35 columnals, sutures flush to depressed; convex aboral surface of brachials. narrows distally; mesistele with nodals and internodals; distis- Description.-Crown narrow, cylindrical. Aboral cup typi- tele not known. cally with perfect pentameral symmetry except for very small Material examined. -The holotype of Eucatillocrinus brad- anal X articulated to upper shoulder of C radial; medium cone leyi is ISGS ISM 2400; and additional material includes FMNH shape; truncate base, and constant to slightly increasing rate of UC 5 1827a, FMNH UC 5 1827b, USNM 14670, and USNM S distal widening; heightwidth ratio ranges from 0.32 to 0.69 22 15 (5 specimens). (mean=0.49, SD=0.07, n=88). Aboral cup plates smooth (Fig- Occurrence. -Eucatillocrinus bradleyi was originally de- ure 5.17) or granulose, very gently convex to very convex, su- scribed from the Edwardsville Formation at Crawfordsville, In- tures flush but suture areas may be depressed (Figure 5.16). diana. We have examined additional specimens from the Ed- Basal plate circlet 14-54 percent of aboral cup height wardsville Formation at Indian Creek, Indiana, in the collec- (mean=32, SD=0.08, n=84); expands in width distally; shallow, tions of Robert Howell. broad basal concavity. Basal plates three, unequal, azygous basal Measurements. -See Table 5. in AE position. Radial circlet height percentage approximately 68; expands distally at same or greater rate than distal expansion Superfamily BELEMNOCRINACEAS. A. Miller, 1883 of basal circlet. Distal to proximal radial plate width ratio ranges Family SYNBATHOCRINIDAES. A. Miller, 1889 from 1.04 to 2.26 (mean= 1.64, SD=0.19, n=88). Radial facet Genus SYNBATHOCRINUSPhillips, 1836 plenary, subhorizontal, narrows adaxially so triangular in shape; Type species. -Synbathocrinus conicus Phillips, 1836; by distally projecting radial processes between adjacent radial fac- monotypy. ets. Radial facet topography includes a long articular ridge that is straight between the lateral abaxial (outside) comers of facet; SYNBATHOCRINUSSWALLOVI Hall, 18 58 an elongate abaxial fossa centrally, abaxial to articular ridge and Figure 5.1-5.2 1 where facet widens centrally (Figure 5.9); triangular adaxial re- gion of facet with a central groove that projects from center of Donacicrinites simplex TROOST,1849, p. 420, (nomen nudum); TROOST, 1850, p. 62, (nomen nudum); SHUMARD,1865, p. 367, (nomen nu- articular ridge adaxially, one fossa on each of the back parts of dum). the facet (total of two), and a series of short ridges that project SynbathocrinusswalloviHw, 1858, p. 672, P1.17, figs. 8,9;WACHSMUTH adaxially from adaxial edge of articular ridge. AND SPRINGER,1886, p. 166 (90), 169 (93); MILLER,1889, p. 285; Very small shoulder on upper margin of C radial in CD in- WELLER,1898, p. 618; BEEDE,1906, p. 1262, P1. 13, fig. 11, P1. 14, terray for very small anal X, which is typically completely above fig. 1; BASSLERANDMOODEY,1943,p. 696;KESLINGANDSM~,1963, the radial circlet. Anal sac relatively short, poorly known. p. 194;VANSANT,1964,p. 71-72, P1. 1, fig. 6,figs. 23.4,26.la-26.le; Arms five, atomous, as much as 24 times higher than aboral STR~MPLEET AL., 197 1, p. 11; WEBSTER,1973, p. 249; AUSICH,1978, cup height (Figure 5.10), broadly convex aborally (Figure 5.21) p. 227; AUSICH,1980, p. 280; AUSICHAND LANE,1980, p. 60, 61; to broadly triangular (Figure 5.10). First primibrachial very dis- AUSICH,1983, p. 257,259, 276; KAMMERETAL.,1983, p. 55, 56, 67; tinctive, approximately same size as radial, well-developed ar- AUSICHAND LANE,1985, p. 220, 224. Synbathocrinw robustus SHUMARD,1866, p. 397;WORTHENAND MEEK, ticular ridge and aboral facet similar to radial plate, gently con- 1875, p. 514, PI. 29, fig. 4;WACHSMUTHANDSPRINGER, 1886, p. 169 vex. Second primibrachial wider than high, subsequent brachi- (93);MILLER,1889, p. 285; WELLER,1898, p. 618; SPRINGER,1923, als gradually changing to approximately as wide as high; deep, P1. 5, figs. 11-12; BASSLERAND MOODEY,1943, p. 696; MOORE AND rounded aboral groove in second and higher primibrachials, LAUWN,1943, P1. 1, figs. 10a, lob; MOOREAND LAUDON,1944, p. ridges may be present on the abaxial lateral margins of the oral 151, P1. 55, fig. 11, P1. 56, fig. 23; KESLINGANDSMITH,1963, p. 194; side of brachials, which interlock with ridges on adjacent bra- WEBSTER,1973, p. 249; WEBSTER,1977, p. 164; MOOREAND LANE, chial~if crown in closed posture. 1978b,p. T559, fig. 352.1c, 352.1d;W~,1982, p. 197,figs.33~- Columnals circular, lumen pentagonal; both nodals and in- 33w; KAMMER,1984, p. 121-122, figs. 2u-2w; WEBSTER,1986, p. 296; WEBSTER,1988, p. 156. ternodals but sequence of internodal addition cannot be deter- Symbathocrinus robustus Shumard, 1866 [sic]. WOOD,1909, p. 28-29, mined with present specimens. Holdfast not known. P1. 4, fig. 11; WOLF,1979, fig. 2a; WILSON,1948, P1. 29, figs. 4, 5. Discussion. -Several specimens (including primary types of Synbathocrinus granuliferus WETHERBY,1880, p. 250-253, P1. 16, figs., two species) assigned here to Synbathocrinus swallovi are part 3, 3a; WACHSMUTHAND SPRINGER,1886, p. 169 (93);MILLER, 1889, of the material described in Troost's unpublished manuscript p. 284, fig. 438; WELLER,1898, p. 617;BASSLERAND MOODEY,1943, (Wood, 1909). In addition to the nomenclatoral confusion sur- p. 695; KESLINGAND SMITH,1963, p. 192; WEBSTER,1973, p. 248. rounding Troost's manuscript (all of Troost's names are nomina Synbathocrinus angularis MILLERAND GURLEY,1894, p. 4243, P1. 4, nuda), Synbathocrinus was among the material Troost mixed figs. 3,4;WELLER,1898, p. 6 17; SPRINGER,191 1, p. 193-194; BASSLER between two localities: Whites Creek Springs, Tennessee (now AND MOODEY,1943, p. 694; -LING AND SMITH,1963, p. 190; WEBSTER,1973, p. 248; KAMMER,1982, p. 197-198, figs. 33r-33t; recognized as the Lower Mississippian Fort Payne Formation) KAMMER, 1984, p. 122, fig. 2r-2t; WEBSTER,1988, p. 155. and Decatur County, Tennessee (now recognized as the Upper Symbathocrinus troosti WOOD,1909 [sic], p. 27, P1. 9, fig. 9. Silurian Brownsport Formation). Wood (1 909) only partially Synbathocrinus troosti (Wood, 1909). BASSLERAND MOODEY,1943, p. clarified this situation, and Bassler and Moodey (1943) finally 696; KESLINGAND SMITH,1963, p. 195; WEBSTER,1973, p. 249. resolved the localities for each species in question. In summary, Synbathocrinites granulatus TROOST,1849 [sic],p. 4 19 (nomen nudum); Troost mixed a Pisocrinus with a Synbathocrinus under the TROOST,1850, p. 61 (nomen nudum). name Synbathocrinus tennesseeae and listed its occurrence as Synbathocrinus granulatus Troost, 1849. SHUMARD,1866, p. 397 (no- both Decatur County and Whites Creek Springs. Roemer (1 860) men nudum); WACHSMUTHAND SPRINGER,1886, p. 166 (nomen nu- was the first to describe this species, and he called it Synbath- dum). Symbathocrinus granulatus WOOD, 1909 [sic], p. 27-28, P1. 9, figs. 4, ocrinus tennesseensis. Wood (1 909) recognized that two species 7. ,-.8 were represented in the Troost material and named a new spe- Synbathocrinus granulatus (Wood, 1909).BASSLERAND MOODEY,1943, cies, S. troosti; but she considered both species to be from the p. 695; KESLINGAND SMITH,1963, p. 192; WEBSTER,1973, p. 248. Silurian of western Tennessee. Wood (1909) was apparently Synbathocrinus sp. MEYER,AUSICH, AND TERRY,1990, fig. 2b. unaware that Bather (1893) had reassigned S. tennesseensis to A USZCH ET AL. -MISSISSIPPIAN DZSPARZD CRZNOZDS 143

FIGURE5-Middle Mississippian synbathombids. 1-21, Synbathocrinus swallovi, 1. lateral view of partial crown of S. swallovi lectotype, UI X-7, x 1.5; 2. 3, FMNH UC 6410 (holotype of S. grandiferm), 2, lateral view of crown, x 1.0, 3, lateral view of aboral cup and proximal-most arms, x 1.5; 4, 5, highly weathered specimen, USNM 39980a (S. troosti syntype), x 1.5, 4, basal view, 5, lateral view, 6, 7, partial specimen, USNM 399801, (S. granulatus syntype), x 2.0,6, basal view, 7, lateral vim, 8, 9, USNM 39980a (S. grandatus syntype), x2.0.8, basal view, 9, lateral view; 10, nearly complete crown, USNM 443601, x 0.6; 11, 12, specimen with straight sides and sutures flush, USNM 483837, x 1.5, 11, basal view, 12, lateral view; 13.14, specimen with depressed sutures, USNM 4838 18, x 1.5.13, basal view, 14, lateral view; 15.16, specimen with deeply depressed sutures, USNM 4838 17, x 1.5.15, basal view, 16, lateral view; 17, lateral view of dorsal cup with well-presewecl radial facets, USNM 483826, x 1.5; 18 lateral view of aboral cup, USNM 399801, (S. troosti syntype), x 1.5; 19, lateral partial crown, FMMI UC 6481 (S. angularis holotype), x 1.5; 20, 21, USNM S 2024b, x 1.5, 20, basal view, 21, lateral view illustrating proximal arms. 22-25, Synbathocrinus blain'. 22.23, lectotype, FMNH UC 6 147b, x 1.522. CD-interray lateral view, 23, A-ray lateral view; 24, lateral view of partial crown with column attached, S. blairi paralectotype, FMMI UC 6147% x 1.5; 25, lateral view of partial, weathered specimen, USNM 483878, x 1.5. JOURNAL OF PALEONTOLOGY, V. 71, NO. 1, 1997

Pisocrinus, but Springer (1 926) established this revised generic assignment. Bassler and Moodey (1943) noted that S. troosti belonged to Synbathocrinus and that it was probably from the Lower Mississippian at Whites Creek Springs, Tennessee. Sim- ilarly, the species now regarded as Synbathocrinus granulatus (Wood) was considered to have been from Decatur County by both Troost and Wood, but again Bassler and Moodey (1943) established that this species was probably from the Lower Mis- sissippian at Whites Creek Springs, Tennessee. Wood (1909) is recognized as the author, herein, because she was the first to describe or illustrate this species. Wood (1909) correctly rec- ognized that Synbathocrinus robustus was initially described by Shumard and it was probably from the Lower Mississippian at

Whites Creek Springs, Tennessee. Our examination of speci- 3 mens agrees with Bassler and Moodey (1943) in that S. troosti, A 1CAN 2 (15%) c - S. granulatus, and S. robustus all belong to Synbathocrinus and PAM all are Lower Mississippian. Similar to Catillocrinus, several species of Synbathocrinus were named from different stratigraphic and geographic posi- tions; but currently there is little rationale to maintain all but two of these species distinctions. Synbathocrinus swallovi, S. robustus, S. granuliferus, S. angularis, S. granulatus, and S. troosti all have a low basal circlet (which distinguishes them FIGURE6-Results from Stepwise Discriminant Function Analysis from S. dentatus from the upper part of the Burlington Lime- (SDFA) of Synbathocrinus,where S. "troosti" is distinguished. CAN1 stone) and smoothly rounded aboral surfaces on brachials. accounts for 85 percent of the variance in the data and is positively Therefore, we regard Synbathocrinus swallovi Hall, 1858 to be correlated with ACH (aboral cup height) and pACW (proximal aboral the senior synonym of these species: Synbathocrinus robustus cup width); CAN2 accounts for 15 percent of the variance in the data Shumard, 1866, Synbathocrinus granuliferus Wetherby, 1880, and is positively correlated with pACW (proximal aboral cup width) Synbathocrinus angularis Miller and Gurley, 1894, Synbatho- and negatively correlated with RRAT (=dRW/pRW, radial plate crinus granulatus (Troost, 1849), and Synbathocrinus troosti shape). The filled circle is S. swallovi holotype, circle with vertical line is S. angularis holotype, filled diamond is S. blairi lectotype, and (Wood, 1909). half-filled diamonds are S. blairi paralectotypes. Originally, S. swallovi was described from the Mississippian type region in Illinois; S. robustus, S. granuliferus, and S. an- gularis were described from the New Providence Shale in Ken- height (ACH), proximal aboral cup width (pACW), distal aboral tucky; and S. granulatus and S. troosti were described from the cup width (dACW), basal circlet height (BCH), distal basal cir- Fort Payne Formation at Whites Creek Spring, Tennessee. Type clet width (dBCW), proximal radial plate width (pRW), and material and/or specimens generally assigned to S. troosti, S. distal radial circlet width (dRW). Ratios for Synbathocrinus are robustus, S. granulatus, and S. angularis tend to be united in height to width ratio ofthe aboral cup (HWRAT=ACH/dACH), sharing quite robust radial plates, separated by a depressed cup relative contribution of the basal circlet to the aboral cup height surface along radial :radial sutures; whereas, S. swallovi and S. (BCPER=BCH/ACH), shape of the radial plates (RRAT=dRW/ granuliferusspecimens have gently convex radial plates and lack pRW), and the basal circlet shape (BCRAT=BCH/dBCW). A a depression along the radia1:radial sutures. Other differences total of 112 specimens were considered in this analysis, Table include size (S. troosti is quite large), plate sculpturing (S.gran- 6 only lists a representative set of specimens. In a SDFA in uliferus), and asymmetries (S. angularis, see Kammer, 1984); which S. troosti, S. swallovi, and S, blairi were chosen as a valid but we do not regard these characters to be substantial or con- species a priori (Figure 6), virtually complete overlap exists sistent enough to warrant species recognition. between S. swallovi and S. troosti, and 20 percent of the spec- The question is whether the S. troosti-type morphology is imens were misclassified (although F statistic probability for distinct from S. swallovi, and we conclude that these two mor- this test is 0.0001). Therefore, the apparent differences in mor- phologies are only end-members of the variation within a single phology cannot be supported morphometrically, and they are species (the senior synonym being S. swallovi Hall, 1858, by considered to be the same species. priority). In large populations the convexity of the radials and In the same SDFA (Figure 6), S. blairi specimens clustered subsequent depth of the radia1:radial depression vary through together with little overlap with S. swallovi. This distinction is the entire morphological range so that only a single population more accurately displayed in an alternate SDFA, comparing is evident. For example, the complete range of morphology is only S. swallovi with S, blairi; and results in virtually non- present from a single population in the Fort Payne Formation overlapping populations with only four percent of specimens of south-central Kentucky, as illustrated in Figure 5.1 1-5.16. misclassified (Figure 7; note that because only two species are Furthermore, other characteristics, such as the arms and calyx compared, only one Canonical Variable is generated). Again, shape, are all similar. The holotype of S. granuliferus is a nearly the F statistic probability for this analysis was 0.000 1. This latter complete crown with the cup somewhat crushed and coarse analysis identified aboral cup height, distal :proximal radial plate preservation of the plate surfaces. Where original plate sculp- width ratio, basal circlet height, and basal circlet percentage turing is preserved, it is clearly within the range present among height of aboral cup as the most important variables, with aboral specimens of S. swallovi. The synonymy of the species is further cup height and basal circlet height relatively more significant supported by the SDFA performed on adult specimens (arbi- than the others. Only very minor overlap occurs between these trarily defined as specimens with an aboral cup height greater two groupings, with only 6.4 percent of the S. swallovi specimens than 3.0 mm). As with other genera, SDFA was performed on misclassified by the test. Synbathocrinus swallovi is differenti- both measurement and ratio data. Measurements are aboral cup ated from S. blairi on the basis of aboral cup size, aboral cup A USICH ET AL. -MISSISSIPPIAN DISPARID CRINOIDS 145

TABLE6-Measurements for Synbathocrinus specimens (in mm). CrH, crown height; ACH, aboral cup height; pACW, proximal aboral cup width; dACW, distal aboral cup width; BCH, basal circlet height; dBCW, distal basal circlet width; pBCW, proximal basal circlet width; pRW, proximal radial plate width; pRW, distal radial plate width. * incomplete or crushed; ** holotype or lectotype. This table includes only select specimens. A complete set of measurements is available from the authors, and a copy is deposited in the Springer Room, U.S. National Museum.

Species/Specimen CrH ACH pACW dACW BCH dBCW PR W dRW

OSU 47694 osu 47696 USNM S 2024a USNM S 2024b USNM S 2024 USNM S 2024 USNM S 2024 USNM S 2024 USNM S 2025 USNM S 2025 USNM S 2025 USNM S 2025 USNM S 2025 USNM S 2031 USNM S 3270 USNM S 3270 USNM S 3270 USNM 14037 USNM 37530 USNM 39938a2 USNM 39938d2 USNM 39980a3 Synbathocrinnus blairi FMNH 1JC 6 147a** FMNH UC 6 I 47b4 FMNH UC 6 147c4 USNM S 2028 USNM S 2029 USNM S 2030 USNM S 2030 USNM S 2030 Holotype of S. angularis. Svntv~esof S, aranulatus. ~oldfypeof S. utroosti. Paralectotypes of S, blairi. shape, aboral plate sculpturing, and the aboral arm surface. Size is unknown. Additional specimens include FMNH UC 8908, is not normally a particularly compelling species character, but ISGS ISM 1830, IU 15144-14008, IU 15124-6, IU 15 172-5064, S. blairi specimens are uniformly smaller than typical specimens IU 15 173-5062, OSU 47692-47696, USNM 14037, USNM of S. swallovi. Accordingly, S. swallovi has larger aboral cups; 37530 (2 specimens), USNM 39940, USNM 42273, USNM commonly medium cone-shaped aboral cups with the sides ei- 443601, USNM 483817-483877, USNM 53270 (3 specimens), ther straight or concave in lateral view; low basal circlet; gently USNM S 2024 (40 specimens), USNM S 2025 (16 specimens), to strongly convex radial plates, radial :radial sutures flush to USNM S 2027, USNM S 2031, USNM S 2062, and USNM S depressed; and convex to somewhat angular aboral surface of 3270. Material previously assigned to S. robustus includes ISGS brachials. In contrast, S. blairi has smaller aboral cups; medium ISM 1830, USNM 2024 (12 specimens), and USNM 39940; bowl aboral cup shape with sides convex; low basal circlet; gently material previously assigned to S. troosti includes USNM convex radial plates, flush radial: radial sutures; and sharply 39980b39980c; and material previously assigned to S. gran- keeled aboral surface of brachials. ulatus includes USNM 39938~-39938d. Material examined. -Hall (1 858) originally illustrated two Occurrence.-Hall (1 858) originally described S. swallovi from specimens, but only a single specimen presently is know to exist. the St. Louis Limestone, St. Louis, Missouri; but labels accom- This specimen (UI X-7) is designated, herein, as the lectotype panying the syntype specimen indicate that Synbathocrinus (Figure 5.1). Type material of junior synonyms include the fol- swallovi was originally described using specimens from the Keo- lowing: Synbathocrinus troosti (Wood, 1909), syntypes, USNM kuk Limestone, Bronsey's Creek, Hancock County, (Hamilton- 39980a (Figure 5.4,s. $5.1 8); Synbathocrinus granulatus (Wood, Warsaw area), Illinois. Junior synonyms were originally de- 1909), two illustrated specimens from the type suite, USNM scribed from the following localities: S. robustus, New Provi- 39938a (Figure 5.6, 5.7) and USNM 39938b (Figure 5.8, 5.9); dence Shale, Button Mold Knob, Kentucky; S. granuliferus, Synbathocrinus granuliferus Wetherby, 1880, holotype, FMNH New Providence Shale, King's Mountain Tunnel along the Cin- UC 6410 (Figure 5.2, 5.3); Synbathocrinus angularis Miller and cinnati Southern Railway, Kentucky; S. angularis, New Prov- Gurley, 1894, holotype, FMNH UC 648 1 (Figure 5.19), and the idence Shale, Button Mold Knob, Kentucky; and S. granulatus location of the types of Synbathocrinus robustus Shumard, 1866, and S. troosti from the Fort Payne Formation, Whites Creek 146 JOURNAL OF PALEONTOLOGY, V. 71, NO. 1, 1997

Diagnosis.-Small aboral cup, medium bowl-shaped ~boral cup with sides convex in lateral view; low basal circlet; gently convex radial plates, radia1:radial sutures flush; sharply keeled aboral surface of brachials. Description. -Crown narrow, cylindrical. Aboral cup typi- cally with perfect pentameral symmetry except for small anal X articulated to upper shoulder of C radial; medium bowl shape; S. blairi truncate base; height: width ratio ranges from 0.33 to 0.60 s. svallori (mean= 0.47, SD=0.05, n=37). Aboral cup plates smooth or granulose, very gently convex to moderately convex (Figure 5.25), sutures flush or suture areas somewhat depressed in some species. Basal plate circlet 16 to 62 percent of aboral cup height (mean=31,SD=0.09, n=33); expands in width distally; shallow, broad basal concavity. Basal plates three, unequal, azygous basal in AE position. Radial circlet height percentage approximately 69. Distal to proximal radial plate width ratio ranges from 1.25 to 1.91 (mean= 1.61, SD=O. 14, n=36). Small shoulder on upper margin of C radial in CD interray

1 1 1 1 for very small anal X, which is typically completely above the ) 1.b 2 .O 3.0 4.0 radial circlet (Figure 5.22,5.24). Anal sac relatively short, poorly CAN 1 known. Arms five, atomous, as much as 20 times higher than aboral FIGURE7-Results from Stepwise Discriminant Function Analysis cup height, angular aborally with distinct ridge (Figure 5.22, (SDFA) of Synbathocrinus where only S, swallovi and S. blairi are 5.23). First primibrachial very distinctive, approximately same distinguished.A histogram illustrating the distribution of CAN1 val- size as radial; very convex with three ridges or folds from each ues for each species is given (CAN 1 accounts for 100 percent of the proximal comer and from center of distal margin. Second pri- variance). mibrachial approximately as high as wide, as are subsequent brachials; deep rounded aboral groove in second and higher Spring, Tennessee. Synbathocrinus swallovi is now recognized primibrachials, ridges may be present on the abaxial lateral from the following localities in the east-central United States: margins of the oral side of brachials that interlock with ridges 1) New Providence Shale Member of the Borden Formation: on adjacent brachials if crown in closed posture. Button Mold Knob, Coral Ridge Quarry, and Kings Tunnel, Columnals circular (Figure 5.24), lumen pentagonal, hetero- Kentucky; and 6 miles west of Jeffersonville, Indiana; 2) Ed- morphic; nodals and internodals alternate in proximal column; wardsville Formation: Crawfordsville, Monroe Reservoir (Al- in mature column nodals separated by one priminternodal, two lens Creek Bank, Boy Scout Camp, Causeway Section, and secundinternodals, and four tertinodals. Holdfast not known. Waldrip Site), Etter Farm (probably Edwardsville Formation), Discussion.-Differentiation of S. blairi from S. swallovi is Stobo bioherm, and Walnut Fork (near Crawfordsville), Indi- principally by the sharp ridges on the aboral surface of brachials, ana; 3) Keokuk Limestone: Green County, Hamilton, Hancock but other differences are discussed above. Because the differ- County, and Nauvoo, Illinois; Iowa Gateway Terminal, Keo- ences in the aboral cups between S. blairi and S. swallovi are kuk, Iowa; 1-55 section, Missouri; 4) Fort Payne Formation: slight and sometimes not possible to determine, it is very prob- Cumberland County (Blacks Ferry, Highway 61 DW, Highway able that S. blairi evolved directly from S. swallovi through 61-S, and Highway 6 1-Ramp), Lake Cumberland (Bugwood cladogenesis. Buildup, Cave Springs North, Cave Springs South, Gross Creek, Material examined.-Syntypes of Synbathocrinus blairi are Harmon Creek Buildup, Owens Branch, Pleasant Hill, Seventy- FMNH UC 6147 (3 specimens). Specimen FMNH UC 6147b Six Falls, Wolf CreekICaney Fork); Celina Buildup, and Whites is designated, herein, as the lectotype, and specimens FMNH Creek Spring, Tennessee; and 5) Lower Warsaw Formation, UC 6147a and FNMH UC 6147c are designated as paralecto- Meramec River Bridge section and Kirkwood, Missouri. types. Additional specimens include USNM 26408 (4 speci- Occurrences listed by Bassler and Moodey (1943, p. 696) from mens), USNM 42277, USNM 212887, USNM 483878, USNM Canton and Spergen Hill, Indiana, and Boonville, Missouri, are S 2028 (16+ specimens), USNM S 2029, USNM S 2030, USNM considered erroneous based upon examination of specimens in S 2031, and USNM S 26408. the Springer Room, USNM. The Salem Limestone occurrence Occurrence.-Synbathocrinus blairi was originally described (Spergen Hill and elsewhere) was apparently based upon isolated from the upper part of the Warsaw Formation at Boonville, aboral cups, making a distinction between S. swallovi and S. Missouri, which is Meramecian in age. Other occurrences are blairi uncertain. Recognition of S. swallovi from the Somerset as follows: 1) Keokuk Limestone; Keokuk, Iowa; 2) Fort Payne Shale Member of the Salem Limestone (Feldman, 1989, p. 908) Formation: Lake Cumberland (Cave Springs South), Kentucky; was also based on isolated aboral cups. 3) Edwardsville Formation: Indian Creek, Indiana; 4) upper part Measurements. -See Table 6. of the Warsaw Formation: Boonville West, Cragwold Road, and Lisbon, Missouri; and Warsaw, Illinois; 5) Harrodsburg Lime- SYNBATHOCRINUSBLAIRI Miller, 1891 stone: Canton, Indiana; 6) Somerset Shale Member of the Salem Figure 5.22-5.25 Limestone: central Kentucky. 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You have printed the following article: Middle Mississippian Disparid Crinoids from the Midcontinental United States William I. Ausich; Thomas W. Kammer; David L. Meyer Journal of Paleontology, Vol. 71, No. 1. (Jan., 1997), pp. 131-148. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28199701%2971%3A1%3C131%3AMMDCFT%3E2.0.CO%3B2-R

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References

A Model for Niche Differentiation in Lower Mississippian Crinoid Communities William I. Ausich Journal of Paleontology, Vol. 54, No. 2. (Mar., 1980), pp. 273-288. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28198003%2954%3A2%3C273%3AAMFNDI%3E2.0.CO%3B2-5

Systematics and Phylogeny of the Late Osagean and Meramecian Crinoids Platycrinites and Eucladocrinus from the Mississippian Stratotype Region William I. Ausich; Thomas W. Kammer Journal of Paleontology, Vol. 64, No. 5. (Sep., 1990), pp. 759-778. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28199009%2964%3A5%3C759%3ASAPOTL%3E2.0.CO%3B2-T

Late Osagean and Meramecian Actinocrinites (Echinodermata: Crinoidea) from the Mississippian Stratotype Region William I. Ausich; Thomas W. Kammer Journal of Paleontology, Vol. 65, No. 3. (May, 1991), pp. 485-499. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28199105%2965%3A3%3C485%3ALOAMA%28%3E2.0.CO%3B2-Y http://www.jstor.org

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Systematic Revisions to Aorocrinus, Dorycrinus, Macrocrinus, Paradichocrinus, Strotocrinus, and Uperocrinus: Mississippian Camerate Crinoids (Echinodermata) from the Stratotype Region William I. Ausich; Thomas W. Kammer Journal of Paleontology, Vol. 65, No. 6. (Nov., 1991), pp. 936-944. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28199111%2965%3A6%3C936%3ASRTADM%3E2.0.CO%3B2-H

Crinoids from the Edwardsville Formation (Lower Mississippian) of Southern Indiana William I. Ausich; N. Gary Lane Journal of Paleontology, Vol. 56, No. 6. (Nov., 1982), pp. 1343-1361. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28198211%2956%3A6%3C1343%3ACFTEF%28%3E2.0.CO%3B2-G

Blastoids from the Late Osagean Fort Payne Formation (Kentucky and Tennessee) William I. Ausich; David L. Meyer Journal of Paleontology, Vol. 62, No. 2. (Mar., 1988), pp. 269-283. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28198803%2962%3A2%3C269%3ABFTLOF%3E2.0.CO%3B2-O

Crinoidea Flexibilia (Echinodermata) from the Fort Payne Formation (Lower Mississippian; Kentucky and Tennessee) William I. Ausich; David L. Meyer Journal of Paleontology, Vol. 66, No. 5. (Sep., 1992), pp. 825-838. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28199209%2966%3A5%3C825%3ACF%28FTF%3E2.0.CO%3B2-V

Fossil Communities of the Borden (Mississippian) Delta in Indiana and Northern Kentucky William I. Ausich; Thomas W. Kammer; N. Gary Lane Journal of Paleontology, Vol. 53, No. 5. (Sep., 1979), pp. 1182-1196. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28197909%2953%3A5%3C1182%3AFCOTB%28%3E2.0.CO%3B2-H http://www.jstor.org

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Demise of the Middle Paleozoic Crinoid Fauna: A Single Extinction Event or Rapid Faunal Turnover? William I. Ausich; Thomas W. Kammer; Tomasz K. Baumiller Paleobiology, Vol. 20, No. 3. (Summer, 1994), pp. 345-361. Stable URL: http://links.jstor.org/sici?sici=0094-8373%28199422%2920%3A3%3C345%3ADOTMPC%3E2.0.CO%3B2-0

Functional Morphology of Calceocrinidae with Description of Some New Species James C. Brower Journal of Paleontology, Vol. 40, No. 3. (May, 1966), pp. 613-634. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28196605%2940%3A3%3C613%3AFMOCWD%3E2.0.CO%3B2-I

The Relations between Allometry, Phylogeny, and Functional Morphology in Some Calceocrinid Crinoids James C. Brower Journal of Paleontology, Vol. 61, No. 5. (Sep., 1987), pp. 999-1032. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28198709%2961%3A5%3C999%3ATRBAPA%3E2.0.CO%3B2-2

Ontogeny and Phylogeny of the Dorsal Cup in Calceocrinid Crinoids James C. Brower Journal of Paleontology, Vol. 64, No. 2. (Mar., 1990), pp. 300-318. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28199003%2964%3A2%3C300%3AOAPOTD%3E2.0.CO%3B2-W

Echinoderms of the Somerset Shale Member, Salem Limestone (Mississippian), in Indiana and Kentucky Howard Randall Feldman Journal of Paleontology, Vol. 63, No. 6. (Nov., 1989), pp. 900-912. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28198911%2963%3A6%3C900%3AEOTSSM%3E2.0.CO%3B2-2 http://www.jstor.org

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Crinoids from the New Providence Shale Member of the Borden Formation (Mississippian) in Kentucky and Indiana Thomas W. Kammer Journal of Paleontology, Vol. 58, No. 1. (Jan., 1984), pp. 115-130. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28198401%2958%3A1%3C115%3ACFTNPS%3E2.0.CO%3B2-D

Aerosol Filtration Theory Applied to Mississippian Deltaic Crinoids Thomas W. Kammer Journal of Paleontology, Vol. 59, No. 3. (May, 1985), pp. 551-560. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28198505%2959%3A3%3C551%3AAFTATM%3E2.0.CO%3B2-1

Aerosol Suspension Feeding and Current Velocities: Distributional Controls for Late Osagean Crinoids Thomas W. Kammer; William I. Ausich Paleobiology, Vol. 13, No. 4. (Autumn, 1987), pp. 379-395. Stable URL: http://links.jstor.org/sici?sici=0094-8373%28198723%2913%3A4%3C379%3AASFACV%3E2.0.CO%3B2-9

Advanced Cladid Crinoids from the Middle Mississippian of the East-Central United States: Intermediate-Grade Calyces Thomas W. Kammer; William I. Ausich Journal of Paleontology, Vol. 67, No. 4. (Jul., 1993), pp. 614-639. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28199307%2967%3A4%3C614%3AACCFTM%3E2.0.CO%3B2-H

Primitive Cladid Crinoids from Upper Osagean-Lower Meramecian (Mississippian) Rocks of East-Central United States Thomas W. Kammer; William I. Ausich Journal of Paleontology, Vol. 70, No. 5. (Sep., 1996), pp. 835-866. Stable URL: http://links.jstor.org/sici?sici=0022-3360%28199609%2970%3A5%3C835%3APCCFUO%3E2.0.CO%3B2-B