209

J. Paleont., 81(1), 2007, pp. 209–212Copyright � 2007, The Paleontological Society0022-3360/07/0081-209$03.00

GILMOCRINUS KENTUCKYENSIS N. SP. FROM THE LATE OSAGEAN(MISSISSIPPIAN) MULDRAUGH MEMBER OF THE BORDEN FORMATION IN

KENTUCKY: A EUROPEAN IMMIGRANT ORIGINALLY DERIVED FROMNORTH AMERICA?

THOMAS W. KAMMER,1 WILLIAM I. AUSICH,2 AND ALAN GOLDSTEIN3

1Department of Geology and Geography, West Virginia University, Morgantown 26506-6300, �tkammer@wvu.edu�, 2Department of GeologicalSciences, 155 South Oval Mall, The Ohio State University, Columbus 43210 �ausich.1@osu.edu�, and 3Falls of the Ohio State Park, 201 West

Riverside Drive, Clarksville, Indiana 47129-3148, �agoldstein@dnr.in.gov�

INTRODUCTION

CRINOIDS FLOURISHED during the Early Mississippian, with lateOsagean crinoids being among the most diverse and well

studied. Among others, late Osagean crinoid faunas include thewell-known Crawfordsville and Indian Creek localities in Mont-gomery County, Indiana (Van Sant and Lane, 1964; Lane, 1973),and the Keokuk Limestone in the Mississippi River valley stra-totype region. The late Osagean faunas in North America havebeen comprehensively revised recently in a series of studies (Au-sich and Kammer, 1990, 1991a, 1991b, 1992; Ausich and Meyer,1992; Kammer and Ausich, 1992, 1993, 1994, 1996; Ausich etal., 1997, 2000; Meyer and Ausich, 1997).

Recently, a new species of the cladid crinoid genus Gilmocri-nus Laudon, 1933 was discovered by Goldstein in the MuldraughMember of the Borden Formation in north-central Kentucky atthe same locality studied by Ausich et al. (2000), raising the totalspecies richness at this locality to 67. This single, small specimenis especially significant because of its implications for the geo-graphic distribution of crinoids between North America and West-ern Europe during the Early Mississippian.

In North America, Gilmocrinus was previously known onlyfrom the late Kinderhookian Stage (Webster, 2003). Gilmocrinuskentuckyensis n. sp. extends the range of the genus into the lateOsagean Stage. There are no reports of Gilmocrinus from theearly and middle Osagean of North America, although there areat least 55 species of advanced cladids known from this timeinterval in the Burlington Limestone (Webster, 2003). In Europe,Gilmocrinus first appears in the early Osagean-equivalent IvorianStage and ranges into the late Osagean-equivalent upper ChadianStage, thus covering the span of time that Gilmocrinus is missingin North America. These biogeographic occurrences suggest thatGilmocrinus originated in North America, migrated to Europe be-fore becoming extinct in North America, and then migrated backto North America before final extinction in the latest Osagean.The paleobiogeography of Gilmocrinus supports the hypothesisof a two-way marine connection between the Eastern Interior Sea-way of North America and Western Europe during the Early Mis-sissippian. In addition to providing a description of a new species,we reassign Ophiurocrinus beggi (Wright, 1938a), from the Isleof Man, to Gilmocrinus.

LOCATION AND STRATIGRAPHY

This new crinoid is from the Muldraugh locality discussed inAusich et al. (2000). It is from a now-flooded and inaccessibleabandoned quarry approximately 3.5 km (2.2 mi) northeast of theI-65 and U.S. Highway 62 and Kentucky Highway 61 interchange(Exit 94 of I-65) (UTM Zone16S, 605,930E, 4,176,920N, WGS84 Elizabethtown 7.5 min topographic quadrangle, Hardin Coun-ty). The quarry was completed in the Muldraugh Member of theBorden Formation and exposed approximately the upper 17 m of

the Muldraugh, as well as the overlying Harrodsburg Limestoneand Somerset Shale Member of the Salem Limestone.

The Muldraugh Member in the Elizabethtown quarry is lateOsagean in age based on conodont biostratigraphy (Nicoll andRexroad, 1975) and the crinoid faunas (Ausich et al., 2000). Au-sich et al. (1994) divided the late Osagean into three time sub-divisions, intervals 3–5. Accordingly, the Muldraugh fauna is cor-relative with the late Osagean interval 4, which includes thewell-known crinoid faunas from the Edwardsville Formation ofMontgomery and Monroe Counties, Indiana (Lane, 1973; Ausichand Lane, 1982); the Fort Payne Formation of south-central Ken-tucky (Ausich and Meyer, 1988, 1990; Meyer and Ausich, 1997);and the upper part of the Keokuk Limestone of the MississippiRiver valley (Kammer et al., 1990). It is time-equivalent with theArundian of Western Europe (Jones, 1996). Additional details ofthe stratigraphy and occurrence of crinoids at this locality are inAusich et al. (2000).

SYSTEMATIC PALEONTOLOGY

Terminology follows Ubaghs (1978), and supergeneric taxon-omy follows Moore and Teichert (1978). This new specimen isreposited in the U.S. National Museum of Natural History(USNM).

Class CRINOIDEA J. S. Miller, 1821Subclass CLADIDA Moore and Laudon, 1943

Order DENDROCRINIDA Bather, 1899Family SCYTALOCRINIDAE Moore and Laudon, 1943

Genus GILMOCRINUS Laudon, 1933Type species.⎯Gilmocrinus iowensis Laudon, 1933; by origi-

nal designation.Diagnosis.⎯High conical cup, infrabasals clearly visible in

side view, peneplenary radial facets, three anal plates in cup, fiveatomous arms with pinnules, prominent anal sac.

DiscussionWe follow McIntosh (2001) in placing pinnulate cladid crinoids

in the order Dendrocrinida, which contains both pinnulate andnonpinnulate taxa. The Poteriocrinida as defined by Moore et al.(1978) is probably a polyphyletic grouping composed of severalpinnulate taxa derived from the Dendrocrinida sensu Moore et al.(1978).

GILMOCRINUS BEGGI (Wright, 1938a) new combinationScytalocrinus beggi WRIGHT, 1938a, p. 285, pl. 11, fig. 12.Gilmocrinus beggi (WRIGHT); WRIGHT, 1938b, p. 343, pl. 13, fig. 2.Ophiurocrinus beggi (WRIGHT); WRIGHT, 1950, p. 22, pl. 6, fig. 6.

Diagnosis.⎯A species of Gilmocrinus lacking interradialnotches and apical pits at the plate corners; brachials mediumlength, subcuneate to cuneate.

Material examined.⎯Holotype and only known specimen isE3354, Hunterian Museum, Glasgow, United Kingdom.

210 JOURNAL OF PALEONTOLOGY, V. 81, NO. 1, 2007

FIGURE 1—Gilmocrinus kentuckyensis n. sp., USNM 528279, holotype; 1, A-ray view of crown, note the very long pinnule along side the A-rayarm, �2.0; 2, A-ray view of aboral cup, �4.0; 3, posterior interarea from back of holotype slab wetted in water, 1 � radianal plate, 2 � anal Xplate, 3 � right tube plate, C � C radial plate, CD � CD basal plate; the apparent hole at the junction of the three anal plates was accidentallyproduced during preparation; �6.0.

TABLE 1—Measurements, in mm, from the anterior of the holotype of Gil-mocrinus kentuckyensis n. sp.

Crown

H

Cup

H W

Infrabasal

H W

Basal

H W

Radial

H W

Pinnule

L

�40.0 5.7 6.4 1.8 1.8 2.6 2.3 2.4 2.9 �11.5

Occurrence.⎯Cass-ny-hawin, Isle of Man, United Kingdom.Wright reports the locality as Visean, C2 of Vaughan (1905),which is equivalent to the upper Chadian (Leeder, 1992).

Discussion.⎯Wright (1950) changed the generic assignmentfrom Gilmocrinus to Ophiurocrinus Jaekel, 1918 because ‘‘thisspecies comes so close to Ophiurocrinus dactyloides (Austin andAustin).’’ Ausich and Sevastopulo (2001) reassigned O. dactyl-oides (Austin and Austin, 1843) to Gilmocrinus. Ophiurocrinusis the incorrect genus because it has plenary radial facets and verylow cuneate brachials with arms nearly circular in cross section.

GILMOCRINUS KENTUCKYENSIS new speciesFigure 1, Table 1

Diagnosis.⎯A species of Gilmocrinus with interradial notches,apical pits where the corners of basals and radials meet, and long,subcuneate brachials.

Description.⎯Crown fairly well preserved, very tall, narrow.Aboral cup cone-shaped, height to width ratio approximately0.80; plates smooth, lacking ornamentation; shallow apical pits

present at corners where basals and radials meet, but absent atinfrabasal-basal corners; infrabasals prominent, clearly visible inside view; basals hexagonal, approximately same size as radials;radials with peneplenary facets and distinct fulcral ridge on thefacets; radials separated by distinct interradial notches. Back ofholotype showing three anal plates in cup in normal arrangementof radianal below and to right of anal X, and right tube platesitting above radianal and to right of anal X; anal sac unknown.Arms atomous in A and B rays, and were presumably in otherrays; brachials high and subcuneate proximally, shorter and morecuneate distally, with extremely long pinnules. Column subpen-tagonal proximally, circular distally, heteromorphic with subtlenodals and internodals.

Etymology.⎯From the state of Kentucky.Type.⎯Holotype and only known specimen is USNM 528279.Measurements.⎯See Table 1.Occurrence.⎯Muldraugh Member of the Borden Formation,

Hardin County, Kentucky, UTM Zone16S, 605,930E,4,176,920N, WGS 84.

Discussion.⎯There are five other known species of Gilmocri-nus: G. iowensis, the type species of the genus from the lateKinderhookian Gilmore City Formation of Iowa; G. oneali Lau-don and Beane, 1937, from the late Kinderhookian Hampton For-mation of Iowa; G. bozemanensis (Miller and Gurley, 1896) fromthe late Kinderhookian Lodgepole Formation of Montana; G. dac-tyloides (Ausich and Sevastopulo, 2001) from the Tournaisian(early Osagean equivalent) Hook Head Formation of Ireland; and

211PALEONTOLOGICAL NOTES

TABLE 2—Stratigraphic and geographic occurrences of species of Gilmocrinus. The Late Osagean spans the upper Chadian and Arundian stages of Europe(Jones, 1996).

Chronostratigraphy North America Europe

Late late Osagean or Arundian G. kentuckyensis n. sp.Early late Osagean or upper Chadian G. beggiMiddle Osagean or lower ChadianEarly Osagean or Ivorian G. dactyloidesLate Kinderhookian or Hastarian G. iowensis, G. oneali, G. bozemanensis

G. beggi from the upper Chadian (late Osagean equivalent) of theIsle of Man. Gilmocrinus kentuckyensis is the youngest speciesof the genus and extends the range into the late Osagean (Table2).

Gilmocrinus kentuckyensis differs from these other species ofGilmocrinus as follows. Gilmocrinus iowensis is similar in manyregards, but lacks distinct interradial notches and apical pits; thebrachials are low cuneate; it also possesses a large anal sac withstellate plates, a feature that cannot be compared with G. ken-tuckyensis because of the lack of preservation of the anal sac.Gilmocrinus oneali also lacks the interradial notches and apicalpits, and in addition has rectangular brachials proximally that be-come subcuneate distally, and the pinnules are extremely short; italso has a hexagonally plated anal tube with no ornamentation.Gilmocrinus bozemanensis is known from one very small speci-men with at least two atomous arms and appears to have thecharacters of Gilmocrinus, although Laudon et al. (1952) incor-rectly placed it in Ramulocrinus Laudon et al., 1952, which hasonly one atomous arm (A ray). Gilmocrinus dactyloides is alsosimilar in many regards, but lacks interradial notches and apicalpits, plus the cup is clearly higher because the basals are moreelongate. Gilmocrinus beggi lacks the distinct interradial notchesand apical pits and has medium length brachials.

Historically, a tendency has existed to assign Lower Mississip-pian crinoids of North America to genera distinct from those inWestern Europe, with primarily only the most common and eu-rytopic taxa being recognized in both regions, such as, for ex-ample, Actinocrinites J. S. Miller, 1821, Platycrinites J. S. Miller,1821, Synbathocrinus Phillips, 1836, and Taxocrinus Phillips inMorris, 1843. This geographic dichotomy was often true for clad-id crinoids. There are important paleobiogeographic distinctionsbetween these two regions, but recent work suggests that LowerMississippian crinoid paleobiogeographic distributions are muchmore complex. For example, Ausich and Sevastopulo (2001) de-scribed Gilmocrinus, Paracosmetocrinus Strimple, 1967, and Sos-tronocrinus Strimple and McGinnis, 1969, from the Ivorian (Tour-naisian) of southern Ireland, and these genera were previouslyknown from only North America. Work in progress by the twosenior authors includes reevaluation of generic assignments ofNorth American and Western European Lower Mississippian cri-noid species for analysis of eurytopy, biogeography, and evolu-tionary paleoecology of these crinoids. Prior to 2001, Gilmocrinuswas only known from the late Kinderhookian of North America.Significantly, new occurrences in Western Europe and the newlate Osagean species reported here provide important new datafor understanding paleobiogeographic controls during this time.

Gilmocrinus kentuckyensis extends the range of the genus intothe late Osagean in North America (Table 2). Previously in NorthAmerica, Gilmocrinus was known only from late Kinderhookianrocks in Iowa and Montana, and it is absent in the early andmiddle Osagean Burlington Limestone or its equivalents. Al-though the absence of Gilmocrinus in these rocks could be dueto collection failure or poor preservation of advanced cladids,there is a significant probability that Gilmocrinus is truly absent.One of us (TWK) has recently documented more than 55 speciesof advanced cladids in the Burlington Limestone and Gilmocrinus

is absent. In Europe, Gilmocrinus occurs in the Ivorian (Tournai-sian Tn3 zone), or early Osagean equivalent, of Ireland (Ausichand Sevastopulo, 2001), and the upper Chadian, or early late Os-agean equivalent, of the Isle of Man, United Kingdom (Wright,1950). Gilmocrinus kentuckyensis is the youngest species of thegenus occurring in the latest Osagean, or Arundian equivalent.

The geographic and temporal distribution of Gilmocrinus sug-gests there might have been a two-way marine connection be-tween the Eastern Interior Seaway of North America and WesternEurope throughout the Early Mississippian. Considering only thisone genus, it is not possible to determine whether biogeographicconnectivity was constant or intermittent. However, current infor-mation suggests that Gilmocrinus originated in North Americaand may have gone extinct on this continent at the end of theKinderhookian but not prior to migration to Europe. It then sur-vived in Europe during the Ivorian and Chadian stages, and waslater reintroduced into North America during the Arundian or lateOsagean equivalent. Of course another possibility is that Gilmo-crinus survived somewhere other than eastern North America orWestern Europe during the Ivorian and Chadian stages beforebeing reintroduced into the Eastern Interior Seaway in the lateOsagean. However, there are no data at present to test this alter-native hypothesis.

ACKNOWLEDGMENTS

This research was supported by the National Science Founda-tion: EAR-0206307 (TWK) and EAR-02059068 (WIA). C. Cookof St. Louis, Missouri, assisted in the preparation of the holotypespecimen. G. D. Webster and G. C. McIntosh provided reviewsthat improved an earlier draft of this manuscript.

REFERENCES

AUSICH, W. I., AND T. W. KAMMER. 1990. Systematics and phylogeny ofthe late Osagean and Meramecian crinoids Platycrinites and Euclado-crinus from the Mississippian stratotype region. Journal of Paleontol-ogy, 64:759–778.

AUSICH, W. I., AND T. W. KAMMER. 1991a. Late Osagean and Merame-cian Actinocrinites from the Mississippian stratotype region (Echino-dermata: Crinoidea). Journal of Paleontology, 65:485–499.

AUSICH, W. I., AND T. W. KAMMER. 1991b. Systematic revisions to Aor-ocrinus, Dorycrinus, Macrocrinus, Paradichocrinus, Strotocrinus, andUperocrinus: Mississippian camerate crinoids from the stratotype re-gion (Echinodermata). Journal of Paleontology, 65:936–944.

AUSICH, W. I., AND T. W. KAMMER. 1992. Dizygocrinus: Mississippiancamerate crinoid from the midcontinental United States (Echinoder-mata). Journal of Paleontology, 66:637–658.

AUSICH, W. I., AND N. G. LANE. 1982. Crinoids from the EdwardsvilleFormation (Lower Mississippian) of Indiana. Journal of Paleontology,56:1343–1361.

AUSICH, W. I., AND D. L. MEYER. 1988. Blastoids from the late OsageanFort Payne Formation (Kentucky and Tennessee). Journal of Paleon-tology, 62:269–283.

AUSICH, W. I., AND D. L. MEYER. 1990. Origin and composition of car-bonate buildups and associated facies in the Fort Payne Formation(Lower Mississippian, south-central Kentucky): An integrated sedi-mentologic and paleontologic analysis. Geological Society of AmericaBulletin, 102:129–146.

212 JOURNAL OF PALEONTOLOGY, V. 81, NO. 1, 2007

AUSICH, W. I., AND D. L. MEYER. 1992. Crinoidea Flexibilia (Echino-dermata) from the Fort Payne Formation (Lower Mississippian; Ken-tucky and Tennessee). Journal of Paleontology, 66:825–838.

AUSICH, W. I., AND G. D. SEVASTOPULO. 2001. The Lower Carboniferous(Tournaisian) crinoids from Hook Head, County Wexford, Ireland. Pa-laeontographical Society Monograph, No. 617, 136 p., 13 pls.

AUSICH, W. I., A. GOLDSTEIN, AND R. YATES. 2000. Crinoids from theMuldraugh Member of the Borden Formation in north-central Ken-tucky (Echinodermata, Lower Mississippian). Journal of Paleontology,74:1072–1082.

AUSICH, W. I., T. W. KAMMER, AND T. K. BAUMILLER. 1994. Demise ofthe Middle Paleozoic crinoid fauna: A single extinction event or rapidfaunal turnover? Paleobiology, 20:345–361.

AUSICH, W. I., T. W. KAMMER, AND D. L. MEYER. 1997. Middle Missis-sippian disparid crinoids from the east-central United States. Journalof Paleontology, 71:131–148.

AUSTIN, T., AND T. AUSTIN JR. 1843. Description of several new generaand species of Crinoidea. Annals and Magazine of Natural History, 11:195–207.

BATHER, F. A. 1899. A phylogenetic classification of the Pelmatozoa.British Association of the Advancement of Science Report, 1898:916–923.

JAEKEL, O. 1918. Phylogenie und System der Pelmatozoen. Palaontolo-gische Zeitschrift, 3:1–128.

JONES, P. J. 1996. Carboniferous (Chart 5), p. 110–126. In G. C. Youngand J. R. Laurie (eds.), An Australian Phanerozoic Timescale. OxfordUniversity Press.

KAMMER, T. W., AND W. I. AUSICH. 1992. Advanced cladid crinoids fromthe middle Mississippian of the east-central United States: Primitive-grade calyces. Journal of Paleontology, 66:461–480.

KAMMER, T. W., AND W. I. AUSICH. 1993. Advanced cladid crinoids fromthe middle Mississippian of the east-central United States: Intermedi-ate-grade calyces. Journal of Paleontology, 67:614–639.

KAMMER, T. W., AND W. I. AUSICH. 1994. Advanced cladid crinoids fromthe middle Mississippian of the east-central United States: Advanced-grade calyces. Journal of Paleontology, 68:339–351.

KAMMER, T. W., AND W. I. AUSICH. 1996. Primitive cladid crinoids fromthe upper Osagean-lower Meramecian (Mississippian) rocks of the east-central United States. Journal of Paleontology, 70:835–866.

KAMMER, T. W., P. L. BRENCKLE, J. L. CARTER, AND W. I. AUSICH. 1990.Redefinition of the Osagean–Meramecian boundary in the Mississip-pian stratotype region. Palaios, 5:414–431.

LANE, N. G. 1973. Paleontology and paleoecology of the Crawfordsvillefossil site (Upper Osagian: Indiana). California University Publicationsin the Geological Sciences, 99, 141 p.

LAUDON, L. R. 1933. The stratigraphy and paleontology of the GilmoreCity Formation of Iowa. University of Iowa Studies, 15(2), 74 p.

LAUDON, L. R., AND B. H. BEANE. 1937. The crinoid fauna of the Hamp-ton Formation at LeGrand, Iowa. University of Iowa Studies, 17(6):226–272.

LAUDON, L. R., J. M. PARKS, A. C. SPRENG. 1952. Mississippian crinoidfauna from the Banff Formation, Sunwapta Pass, Alberta. Journal ofPaleontology, 26:544–575.

LEEDER, M. R. 1992. Dinantian, p. 207–238. In P. McL. D. Duff and A.J. Smith (eds.), Geology of England and Wales. Geological Society ofLondon.

MCINTOSH, G. C., 2001. Devonian cladid crinoids: Families Glossocrin-idae Goldring, 1923, and Rutkowskicrinidae new family. Journal ofPaleontology, 75:783–807.

MEYER, D. L., AND W. I. AUSICH. 1997. Morphologic variation withinand between populations of the camerate crinoid Agaricocrinus (LowerMississippian, Kentucky and Tennessee): Breaking the spell of themushroom. Journal of Paleontology, 71:896–917.

MILLER, J. S. 1821. A Natural History of the Crinoidea or Lily-ShapedAnimals, With Observations on the Genera Asteria, Euryale, Comatula,and Marsupites. Bryan, Bristol, 150 p.

MILLER, S. A., AND W. F. E. GURLEY. 1896. New species of Echinoder-mata and a new crustacean from the Palaeozoic rocks. Illinois StateMuseum of Natural History Bulletin, 10, 91 p.

MOORE, R. C., AND L. R. LAUDON. 1943. Evolution and classification ofPaleozoic crinoids. Geological Society of America Special Paper, 46,153 p.

MOORE, R. C., AND C. TEICHERT (EDS.). 1978. Treatise on InvertebratePaleontology. Pt. T. Echinodermata 2. Geological Society of Americaand University of Kansas Press, Lawrence.

MOORE, R. C., N. G. LANE, AND H. L. STRIMPLE. 1978. Order Cladida,p. T578–T755. In R. C. Moore and C. Teichert (eds.), Treatise onInvertebrate Paleontology. Pt. T. Echinodermata 2(2). Geological So-ciety of America and University of Kansas Press, Lawrence.

MORRIS, J. 1843. A Catalogue of British Fossils. Comprising all the gen-era and species hitherto described; with reference to the geologicaldistribution and to the localities in which they have been found. JohnVan Voorst, London, 222 p.

NICOLL, R. S., AND C. B. REXROAD. 1975. Stratigraphy and conodontpaleontology of the Sanders Group (Mississippian) in Indiana and ad-jacent Kentucky. Indiana Geological Survey Bulletin, 51, 36 p.

PHILLIPS, J. 1836. Illustrations of the Geology of Yorkshire, or A De-scription of the Strata and Organic Remains. The Mountain LimestoneDistricts, Pt. 2 (second edition). London, John Murray, p. 203–208.

STRIMPLE, H. L. 1967. Aphelecrinidae, a new family of inadunate cri-noids. Okalahoma Geology Notes, 28:33–36.

STRIMPLE, H. L., AND M. R. MCGINNIS. 1969. New crinoid from theGilmore City Formation, Lower Mississippian of Iowa. Fossil CrinoidStudies, University of Kansas Paleontological Contributions, Paper42(5):21–22.

UBAGHS, G. 1978. Skeletal morphology of fossil crinoids, p. T58–T216.In R. C. Moore and C. Teichert (eds.), Treatise on Invertebrate Pale-ontology. Pt. T. Echinodermata 2(1). Geological Society of Americaand University of Kansas Press, Lawrence.

VAN SANT, J. F., AND N. G. LANE. 1964. Crawfordsville crinoid studies.University of Kansas Paleontological Contributions, Article 7, 136 p.

VAUGHAN, A. 1905. The palaeontological sequence in the CarboniferousLimestone of the Bristol area. Quarterly Journal of the Geological So-ciety of London, 61:181–307.

WEBSTER, G. D. 2003. Bibliography and Index of Paleozoic Crinoids,Coronates, and Hemistreptocrinoids, 1758–1999. Geological Society ofAmerica Special Paper, 363 (CD). URL: crinoid.gsajournals.org/crinoidmod

WRIGHT, J. 1938a. Some British Platycrinidae and descriptions of newspecies. Geological Magazine, 75:266–287.

WRIGHT, J. 1938b. Anemetocrinus N.G., a five-armed poteriocrinid fromthe Lower Carboniferous limestones of Scotland. Geological Magazine,75:337–346.

WRIGHT, J. 1950. The British Carboniferous Crinoidea. Palaeontograph-ical Society Monograph, 1(1):1–24.

ACCEPTED 5 SEPTEMBER 2005