A Revision of the Turonian Members of the Ammonite Subfamily Collignoniceratinae from the United States Western Interior and Gulf Coast

Home , Acanthoceratinae, Barroisiceras, Collignoniceras, Collignoniceratidae, Watinoceras

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_1 Allen Press • DTPro System File # 01235TQ

A REVISION OF THE TURONIAN MEMBERS OF THE AMMONITE SUBFAMILY COLLIGNONICERATINAE FROM THE UNITED STATES WESTERN INTERIOR AND GULF COAST

W. JAMES KENNEDY Curator, Geological Collections University Museum of Natural History Parks Road, Oxford OX1 3PW, United Kingdom e-mail: [email protected]

WILLIAM A. COBBAN Research Associate, Division of Paleontology (Invertebrates) American Museum of Natural History Home address: 70 Estes St., Lakewood, CO 80226

NEIL H. LANDMAN Curator, Division of Paleontology (Invertebrates) American Museum of Natural History e-mail: [email protected]

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 267, 148 pp., 124 ®gures, 13 tables Issued December 13, 2001

Copyright ᭧ American Museum of Natural History 2001 ISSN 0003-0090 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_2 Allen Press • DTPro System File # 01235TQ

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CONTENTS Abstract ...... 3 Introduction ...... 3 Localities of Illustrated Fossils ...... 4 Biostratigraphic Distribution of the Collignoniceratinae ...... 13 Variation Within the Collignoniceratinae ...... 27 Conventions ...... 29 Systematic Paleontology ...... 29 Family COLLIGNONICERATIDAE Wright and Wright, 1951 ...... 29 Subfamily COLLIGNONICERATINAE Wright and Wright, 1951 ...... 29 Genus Cibolaites Cobban and Hook, 1983 ...... 29 Cibolaites molenaari Cobban and Hook, 1983 ...... 34 Genus Collignoniceras Breistroffer, 1947 ...... 41 Collignoniceras woollgari woollgari (Mantell, 1822) ...... 42 Collignoniceras woollgari regulare (Haas, 1946) ...... 45 Collignoniceras vermilionense (Meek and Hayden, 1860) ...... 57 Collignoniceras jorgenseni, new species ...... 61 Collignoniceras praecox (Haas, 1946) ...... 65 Collignoniceras percarinatum (Hall and Meek, 1856) ...... 72 Genus Collignonicerites, new genus ...... 76 Collignonicerites collisniger, new genus, new species ...... 81 Genus Prionocyclus Meek, 1876 ...... 81 Prionocyclus hyatti (Stanton, 1894) ...... 83 Prionocyclus albinus (Fritsch, 1872) ...... 93 Prionocyclus macombi Meek, 1876 ...... 95 Prionocyclus bosquensis Kennedy, 1988 ...... 105 Prionocyclus wyomingensis Meek, 1876 ...... 109 Prionocyclus novimexicanus (Marcou, 1858) ...... 117 Prionocyclus quadratus Cobban, 1953 ...... 121 Prionocyclus germari (Reuss, 1845) ...... 123 Prionocyclus pluricostatus, new species ...... 127 Genus Prionocyclites Kennedy, 1988 ...... 129 Prionocyclites mite Kennedy, 1988 ...... 129 Genus Reesidites Wright and Matsumoto, 1954 ...... 139 Reesidites minimus (Hayasaka and Fukada, 1951) ...... 139 Acknowledgments ...... 140 References ...... 141 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_3 Allen Press • DTPro System File # 01235TQ

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ABSTRACT The Collignoniceratinae of the U.S. Western Interior ®rst appear in the lower Turonian and range through the middle, and most of the upper, Turonian. For most of their range they are important biostratigraphic indicators. A review of their biostratigraphic distribution is provided and this is followed by a revision and/or discussion of the following taxa: Cibolaites molenaari Cobban and Hook, 1983, Collignoniceras woollgari woollgari (Mantell, 1822), C. woollgari regulare Haas, 1946, C. vermilionense (Meek and Hayden, 1860), C. jorgenseni n. sp., C. percarinatum (Hall and Meek, 1856), C. praecox (Haas, 1946), Collignonicerites collisniger n. gen., n. sp., Prionocyclus hyatti (Stanton, 1894), P. albinus (Fritsch, 1872), P. macombi Meek, 1876, P. bosquensis Kennedy, 1988, P. wyomingensis Meek, 1876, P. novimexicanus (Marcou, 1858), P. quadratus Cobban, 1953, P. germari (Reuss, 1845), P. pluricostatus n. sp., Prionocyclites mite Kennedy, 1988, and Reesidites minimus Hayasaka and Fukada, 1951. Collignoniceras woollgari, Prionocyclus hyatti, P. macombi, P. wyomingensis, P. novimexicanus, P. quadratus, and P. germari have been designated as zonal species in many publications, but aside from C. woollgari, P. hyatti, and P. wyomingensis, most of these species have not been well described in terms of their intraspeci®c variation. In many species, there is a gracile and a robust form, which may represent sexual dimorphs, but in most instances, the sample size of adult specimens is too small to prove this.

INTRODUCTION Members of the subfamily were collected and described during the early surveys of the The Collignoniceratidae are a family of United States Interior, notably by Marcou Late Cretaceous (latest Cenomanian to mid- (1858), Hall and Meek (1856), Meek (1870, dle Campanian) ammonites with a wide- 1871, 1876a, 1876b), and Stanton (1894), spread distribution (North and South Amer- while material from the Gulf Coast was de- ica, Europe, Africa, and Asia). The species scribed by Shumard (1860). Moreman (1927, are characterized by their involute to evolute 1942) redescribed several species from coiling, generally compressed whorls, and north-central Texas, but illustrated only lim- oval to rectangular whorl section. The venter ited material, and Adkins (1931) and Sidwell commonly bears one or several entire or ser- (1932) described specimens from Trans-Pe- rated keels and the ¯anks and ventrolateral shoulders are ornamented by as many as six cos Texas and the Western Interior, respec- rows of tubercles. The Subfamily Collignon- tively. Haas (1946) provided a detailed ac- iceratinae appeared in the latest Cenomanian count of variation in ``Prionotropis'' wooll- and persisted into the Coniacian, reaching its gari and Prionocyclus wyomingensis from maximum diversity in the Turonian. Many the northern part of the Western Interior, but, species such as Collignoniceras woollgari unfortunately, most of his material lacked (Mantell, 1822), Prionocyclus hyatti (Stan- precise stratigraphic and geographic data. ton, 1894), P. germari (Reuss, 1845), P. al- Powell (1963) described material from north- binus (Fritsch, 1872), and Cibolaites mole- central Texas and Chihuahua, Mexico. naari (Cobban and Hook, 1983) are impor- Matsumoto (1965, 1971) discussed the tant biostratigraphic markers (Matsumoto, genera and certain species of Collignonicer- 1959, 1965, 1977; Wright, 1979, 1996; atinae from North America in his revision of AmeÂdro and Badillet, 1982; Cobban, 1984a; the Japanese members of the subfamily. Cob- Kaplan, 1988; Hancock et al., 1993; Marci- ban (1953) described the new species Prion- nowski et al., 1996). ocyclus quadratus from the Interior, and re- We describe the representatives of the sub- viewed other species with colleagues in subfamily that occur in the Western Interior and sequent publications (Cobban, 1983; Cobban Gulf Coast regions of the United States (®g. et al., 1956; Cobban and Scott, 1972; Cobban 1), a region where collignoniceratines are and Hook, 1979, 1983; Hook and Cobban, more abundant than in any other area yet de- 1979, 1980). More recently, Kennedy and scribed, especially in the middle and upper Cobban (1988), Kennedy et al. (1989), and Turonian. Kennedy (1988) described material from BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_4 Allen Press • DTPro System File # 01235TQ

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Chihuahua, Mexico, Trans-Pecos Texas, and of Fairburn, Custer County, South Dakota. northeast Texas, respectively. These recent Carlile Shale, from a limestone concretion in accounts, plus revisions of the type material the lower part. of the European species (Kennedy et al., 12. D13832. Sec. 35, T. 8 S, R. 1 E, Fall 1980; Wright and Kennedy, 1981; Kaplan, River County, South Dakota. Carlile Shale, 1988) provide a framework in which to de- from limestone concretions in lower part. scribe the many thousands of specimens in 13. D8399. Sec. 2, T. 9 S, R. 1 E, Fall the collections of the U. S. Geological Sur- River County, South Dakota. Carlile Shale, vey and place these specimens in a chron- Turner Sandy Member. ostratigraphic sequence in order to establish 14. D13833. NW¼ SW¼ sec. 8, T. 9 S, R. an international faunal timescale. 2 E, Fall River County, South Dakota. Carlile Shale, from limestone concretions in lower LOCALITIES OF ILLUSTRATED FOSSILS part of Pool Creek Member. Figure 1 15. D10697. SE¼ SE¼ sec. 18, T. 9 S, R. 2 E, Fall River County, South Dakota. Carlile 1. 21191. North of Belle Fourche in the Shale, from limestone concretions about 6 m N½ sec. 10, T. 9 N, R. 2 E, Butte County, (20 ft) below base of Turner Sandy Member. South Dakota. Carlile Shale, from concre- 16. D13834. SW¼ SW¼ sec. 25, T. 7 S, tions 72±73 m (235±240 ft) above base. R. 6 E, Fall River County, South Dakota. 2. 21192. Same locality as 21191. Carlile Carlile Shale, from a limestone concretion in Shale, from concretions 73±76 m (240±250 lower part. ft) above base. 17. D13185. NE¼ SW¼ sec. 26, T. 7 S, 3. 21194. Same locality as 21191. Carlile R. 6 E, Fall River County, South Dakota. Shale, from concretions 76.5±80.4 m (251± Carlile Shale, from white limestone concre- 264 ft) above base. tions in Pool Creek Member. 4. 21195. North of Belle Fourche in the 18. D12215. Angostura Reservoir in the NW¼ sec. 10, T. 9 N, R. 2 E, Butte County, W½ sec. 34, T. 8 S, R. 6 E, Fall River Coun- South Dakota. Carlile Shale, from concre- ty, South Dakota. Carlile Shale, from sand- tions 83±83.5 m (272±274 ft) above base. stone concretions 17 m above base of Turner 5. 21183. North of Belle Fourche in the Sandy Member. SW¼ sec. 11, T. 9 N, R. 2 E, Butte County, 19. D203. NE¼ NW¼ sec. 28, T. 35 N, R. South Dakota. Carlile Shale, from concre- 47 W, Dawes County, Nebraska. Carlile tions 17.7±19.8 m (58±65 ft) above base. Shale, from large laminated silty limestone 6. 21187. Same locality as 21183. Carlile concretions 40±40.5 m (131±133 ft) above Shale, from concretions 34±35 m (111±114 base. ft) above base. 20. D12683. Old dam site on Brule Creek 7. D9896. NW¼ sec. 35, T. 46 N, R. 63 in the NE¼ NW¼ sec. 32, T. 94 N, R. 50 W, W, Weston County, Wyoming. Carlile Shale, Union County State Park, Union County, from limestone concretions 18.3 m below South Dakota. Carlile Shale. base of Turner Sandy Member. 21. D10404. Ionia Volcano in the NW¼ 8. 21792. West of Newcastle in the NW¼ SE¼ sec. 3, T. 31 N, R. 5 E, Dixon County, sec. 31, T. 45 N, R. 61 W, Weston County, Nebraska. Carlile Shale. Wyoming. Carlile Shale, from limestone 22. D1836. North Solomon River in the concretions 18.3 m (60 ft) below base of NE¼ sec. 27, T. 5 S, R. 13 W, Smith County, Turner Sandy Member. Kansas. Carlile Shale, from ¯at ferruginous 9. D8849. Sec. 31, T. 37 N, R 61 W, Ni- concretions in lower part of Blue Hill Mem- obrara County, Wyoming. Carlile Shale, ber. from middle of Turner Sandy Member. 23. 21838. About 4.8 km (3 mi) south- 10. D8443. Sec. 12, T. 36 N, R. 62 W, southeast of Tipton in the SE¼ sec. 4, T. 9 Niobrara County, Wyoming. Carlile Shale, S, R. 10 W, Mitchell County, Kansas. Carlile 16 m (52.5 ft) above base of Turner Sandy Shale, from limestone concretions in upper Member. part of Blue Hill Member. 11. 18872. About 3.2 km (2 mi) southeast 24. D9833. NW¼ NW¼ sec. 33, T. 40 N, BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_5 Allen Press • DTPro System File # 01235TQ

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Fig. 1. Map of the U.S. Western Interior showing the localities of fossils described in the present study. See text for a description of the localities. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_6 Allen Press • DTPro System File # 01235TQ

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Fig. 2. Turonian ammonite zonation used in the text, based on Cobban (1984a), Kennedy et al. (1989), Hancock et al. (1993), Cobban (1990), and Walaszczyk and Cobban (2000).

R. 82 W, Natrona County, Wyoming. Fron- sec. 18, T. 39 N, R. 82 W, Natrona County, tier Formation, from second ledge-forming Wyoming. Frontier Formation, Wall Creek sandstone below top. Member. 25. D12956. At the Gap in the NE¼ NE¼ 26. D9118. NW¼ NE¼ sec. 4, T. 33 N, R. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_7 Allen Press • DTPro System File # 01235TQ

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Fig. 3. Stratigraphic distribution of Collignoniceratinae in the U.S. Western Interior.

81 W, Natrona County, Wyoming. Frontier Formation, from small limestone concretions Formation, from uppermost sandstone of 7±9 m above a sandstone unit bearing Col- Wall Creek Member. lignoniceras woollgari. 27. D9877. SE¼ NE¼ sec. 34, T. 33 N, R. 28. D3763. About 2 km north of Medicine 82 W, Natrona County, Wyoming. Frontier Bow in center of the SE¼ sec. 32, T. 23 N, BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_8 Allen Press • DTPro System File # 01235TQ

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Fig. 4. Measurements used in the text: D ϭ diameter, Wb ϭ whorl breadth, Wh ϭ whorl height, U ϭ umbilical diameter.

R. 78 W, Carbon County, Wyoming. Frontier 39. D4395. North of Rio Gallina in SE¼ Formation, from basal ledges of Wall Creek sec. 15, T. 25 N, R. 1 E, Rio Arriba County, Member. New Mexico. Mancos Shale, from base of 29. D9244. NE¼ sec. 7, T. 21 N, R. 74 W, Juana Lopez Member. Albany County, Wyoming. Frontier Forma- 40. D3884. Arroyo Lopez 1 km north of tion, from lowest ledge-forming sandstone Holy Ghost Spring, Sandoval County, New bed of Wall Creek Member. Mexico. Mancos Shale, from basal part of 30. D6928. NE¼ SW¼ sec. 31, T. 22 N, Semilla Sandstone Member (for map, see R. 75 W, Albany County, Wyoming. Frontier Dane et al., 1968). Formation, Wall Creek Sandstone Member. 41. 28873. Arroyo Lopez 0.6±0.9 km 31. D8981. SE¼ NW¼ sec. 25, T. 19 N, north of Holy Ghost Spring, Sandoval Coun- R. 71 W, Albany County, Wyoming. Frontier ty, New Mexico. Mancos Shale, from basal Formation, from uppermost part of Wall part of Semilla Sandstone Member. Creek Sandstone Member. 42. D3694. About 13.3 km (8.3 mi) south 32. 23422. SE¼ sec. 15, T. 15 S, R. 12 E, of La Ventana, Sandoval County, New Mex- Carbon County, Utah. Mancos Shale, from ico. Mancos Shale, from basal part of Sem- silty, septarian, limestone concretions below illa Sandstone Member. the Ferron Sandstone Member. 43. D5349. About 3.2 km (2 mi) east of 33. D7227. About 8 km east of Ferron, in Seboyeta, Cibola County, New Mexico. the NW¼ sec. 9, T. 20 S, R. 8 E, Emery Mancos Shale, from a limestone concretion County, Utah. Mancos Shale, about 30 m (98 18 m (60 ft) above top of main ledge of Ju- ft) above Ferron Sandstone Member. ana Lopez Member. 34. D11898. NW¼ SE¼ sec. 22, T. 8 S, 44. D11208. NE¼ sec. 36, T. 6 N, R. 19 R. 87 W, Pitkin County, Colorado. Mancos W, Cibola County, New Mexico. Mancos Shale, from calcareous shale 4 m (13 ft) be- Shale, D-Cross Member. low base of Fort Hays Limestone Member. 45. D11281. NE¼ sec. 2, T. 5 N, R. 20 W, 35. D3993. NW¼ sec. 20, T. 20 S, R. 65 Cibola County, New Mexico. Same strati- W, Pueblo County, Colorado. Carlile Shale, graphic position as D8429. from concretion in Blue Hill Member. 46. D11342. NE¼ sec. 2, T. 5 N, R. 19 W, 36. 17632. North of El Vado, Rio Arriba Cibola County, New Mexico. Mancos Shale, County, New Mexico. Mancos Shale, Juana near top. Lopez Member. 47. D8429. Sec. 1 and NE¼ sec. 12, T. 4 37. D2895. About 14 km south of Chama, N, R. 19 W, Cibola County, New Mexico. Rio Arriba County, New Mexico. Mancos Mancos Shale, from limestone concretions Shale, Juana Lopez Member. 24±30 m (78±97 ft) below top of Rio Salado 38. D4407. NW¼ sec. 25, T. 26 N, R. 4 Tongue. E, Rio Arriba County, New Mexico. Mancos 48. D10525. W½ NW¼ sec. 10, T. 4 N, R. Shale, Juana Lopez Member. 7 W, Cibola County, New Mexico. Mancos BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_9 Allen Press • DTPro System File # 01235TQ

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Fig. 5. Cibolaites molenaari Cobban and Hook, 1983. Holotype, USNM 328766, a gracile individual from locality 44. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_10 Allen Press • DTPro System File # 01235TQ

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Fig. 6. Cibolaites molenaari Cobban and Hook, 1983. A±C. Paratype USNM 328761, a gracile BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_11 Allen Press • DTPro System File # 01235TQ

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TABLE 1 Dimensions (mm) of Cibolaites molenaari Cobban and Hook, 1983a

Shale, from septarian limestone concretions E, Socorro County, New Mexico. Mancos in Rio Salado Tongue. Shale, from Rio Salado Tongue. 49. D5773. NW¼ NW¼ sec. 24, T. 3 N, 54. D10643. SE¼ SW¼ sec. 24, T. 9 S, R. R. 7 W, Socorro County, New Mexico. Man- 8 E, Lincoln County, New Mexico. Mancos cos Shale, from Rio Salado Tongue. Shale, from sandy limestone concretions in 50. D10297. NW¼ NW¼ sec. 29, T. 3 N, Rio Salado Tongue. R. 8 W, Socorro County, New Mexico. Top 55. D10644. SW¼ SE¼ sec. 24, T. 9 S, R. of Rio Salado Tongue of Mancos Shale 8 E, Lincoln County, New Mexico. Mancos (Hook et al., 1983: sheet 1). Shale, from sandstone concretion 9 m (30 ft) 51. D10127. N½ sec. 7, T. 2 N, R. 5 W, above base of D-Cross Tongue. Socorro County, New Mexico. Mancos 56. D10636. NE¼ NE¼ sec. 25, T. 9 S, R. Shale, from a little above Juana Lopez Mem- 8 E, Lincoln County, New Mexico. D-Cross ber. Tongue of Mancos Shale, from sandstone 52. D10240. SE¼ SE¼ sec. 8, T. 5 S, R. concretions 9 m (29.5 ft) above base. 2 E, Socorro County, New Mexico. Rio Sal- 57. D10120. East side of Mescal Canyon ado Tongue of Mancos Shale. about 1 km above mouth, Sierra County, New 53. D10243. E½ NE¼ sec. 9, T. 5 S, R. 2 Mexico. D-Cross Tongue of Mancos Shale.

← individual from locality 46. D. Paratype USNM 328765, a robust individual from locality 47. E±G. Paratype USNM 328757, a gracile individual from locality 45. H±J. Paratype USNM 328762, a gracile individual from locality 47. K±M. USNM 498202, a robust individual from locality 45. N±P. USNM 328763, a gracile form from locality 47. Q±S. USNM 498203, a robust form from locality 44. T, U. Paratype USNM 328755, a gracile form from locality 47. V, W. USNM 498212, a robust form from locality 47. X, Y. USNM 498205, a robust form from locality 47. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_12 Allen Press • DTPro System File # 01235TQ

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58. D2495. SW¼ sec. 19, T. 20 S, R. 4 E, DonÄa Ana County, New Mexico. Mancos Shale, from concretions in D-Cross Member. 59. D12671. North of Davis Well in the NW¼ NW¼ sec. 19, T. 20 S, R. 4 E, DonÄa Ana County, New Mexico. Mancos Shale, from limestone concretions in D-Cross Tongue. Fig. 8. Cibolaites molenaari Cobban and 60. D2494. SW sec. 19, T. 20 S, R. 4 E, Hook, 1983. External suture line of USNM ¼ 328760 from locality 44. From Cobban and Hook DonÄa Ana County, New Mexico. Mancos (1983, ®g. 14C). Shale, from concretions in D-Cross Member. 61. D12672. SW¼ sec. 30, T. 20 S, R. 4 E, DonÄa Ana County, New Mexico. D-Cross Tongue of Mancos Shale. (1978). They regarded the highest occurrence of the ammonite Sciponoceras gracile 62. D11186. One km N 87ЊE of Needle Peak, Jeff Davis County, Texas. Chispa Sum- as the base of the Turonian, and the appear- mit Formation. ance of Collignoniceras and Prionocyclus 63. 22608. White Rock Scarp 3.2 km west as the base of the middle Turonian. A re- of Cedar Hill, Dallas County, Texas. Arcadia gional disconformity marked the base of Park Formation, 23 m (75 ft) below top. rocks assigned to the upper Turonian. The base of the upper Turonian was more for- mally placed at the ®rst appearance of the BIOSTRATIGRAPHIC DISTRIBUTION ammonite Prionocyclus macombi by Cob- OF THE COLLIGNONICERATINAE ban and Hook (1979), but Walaszczyk and The ammonite zones recognized here are Cobban (2000) have recently shown that it shown in Figure 2, and are based on the work occurs at a higher position (base of Scaphi- of Cobban (1984a), Kennedy et al. (1989), tes whit®eldi Zone). Hancock et al. (1993), Cobban (1990), and Figure 3 summarizes the biostratigraphic Walaszczyk and Cobban (2000), where de- range distribution of the subfamily in the tailed supporting arguments for the succes- U.S. Western Interior. The earliest collignon- sion are set out. iceratine is Cibolaites Cobban and Hook, In recent years, the Turonian Stage has 1983, with type and only described species been informally subdivided into lower, mid- Cibolaites molenaari Cobban and Hook, dle, and upper Turonian substages. During 1983 (p. 16, pl. 2, ®gs. 1±9; pl. 3, ®gs. 3±8; the 1995 Second International Symposium pl. 8, ®gs. 6±8; pl. 13, ®gs. 1±6; text-®gs. on Cretaceous Stage Boundaries in Brussels 13, 14). This was originally described from (Rawson et al., 1996), the base of the lower the upper lower Turonian Mammites nodo- Turonian was placed at the ®rst occurrence soides Zone of New Mexico, but the genus of the ammonite Watinoceras devonense, and has subsequently been reported (as C. cf. mo- the base of the middle Turonian was placed lenaari) from the upper Cenomanian Neo- at the lowest occurrence of the ammonite cardioceras juddii Zone of Aube, France Collignoniceras woollgari (see Bengston, (Kennedy et al., 1986: 209, ®g. 5i, j, k, l) 1996). No proposal was set forth for the base and the lower Turonian of Germany (Kaplan, of the upper Turonian. 1988). Turonian rocks in the Western Interior re- Collignoniceras? sp. of Wright and Ken- gion were ®rst assigned to lower, middle, nedy (1981: 107, pl. 8, ®g. 17) from the up- and upper Turonian by Kauffman et al. per Cenomanian of Devon, England, also ap-

← Fig. 7. Cibolaites molenaari Cobban and Hook, 1983. A. USNM 498212; B. USNM 498213, both robust forms from locality 47. C, D. USNM 498214; E, H. USNM 498206; F, G. USNM 356901; I. USNM 498207, all robust forms from locality 44. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_14 Allen Press • DTPro System File # 01235TQ

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Fig. 9. Lecointriceras ¯euriausianum (d'Orbigny, 1841). A±C. No. 629b in the Collections of the ChaÃteau de Saumur, from the lower middle Turonian in the environs of Saumur, Maine-et-Loire, France. D, E. No. 122 in the Collections of the FaculteÂ des Sciences, Le Mans, from the same horizon in Sarthe, France. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_15 Allen Press • DTPro System File # 01235TQ

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Fig. 10. Collignoniceras woollgari woollgari (Mantell, 1822). The lectotype BMNH 5682, the original of Mantell, 1822: plate 21, ®gure 6, from near Lewes, Sussex, England. Figures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_16 Allen Press • DTPro System File # 01235TQ

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pears to be a Cibolaites. The distinctive fea- as a distinctive body chamber ornament, it tures of the genus are the presence, during more closely resembles the morphology of most of ontogeny, of umbilical, ventrolateral, stratigraphically younger Collignoniceras and siphonal tubercles. It differs from most species. contemporaneous Acanthoceratinae in that it Collignoniceras ranges through the wooll- only has a single row of ventrolateral tuber- gari Zone and into the neptuni Zone in Eu- cles. Apart from this difference, the gross rope (Kennedy et al., 1986); in the U.S. shell form of certain Thomelites Wright and Western Interior it disappears somewhat ear- Kennedy, 1973 (in Juignet et al., 1973) is lier, prior to the Prionocyclus hyatti Zone. similar to Cibolaites (Wright and Kennedy, The type species, C. woollgari (Mantell, 1990: 285, pls. 80±84). 1822), occurs in both regions as well as in Cibolaites is succeeded stratigraphically in Turkmenistan, Japan, California, Oregon, Europe by Lecointriceras Kennedy et al., and northern Australia. Species of the genus 1980 (type species Lecointriceras ¯euriau- are dimorphic (Kennedy et al., 1989), with sianum (D'Orbigny, 1841) (p. 350, pl. 107, differences in size and ornament. ®gs. 1±3)), a genus widespread in Touraine Collignoniceras woollgari can be distin- and Aquitaine in France but also recorded guished from Lecointriceras in that the latter from northern Spain, the Czech Republic, has a trapezoidal whorl section, and sparse north Germany, and southern England. The low ribs in early growth, with blunt ventro- genus is middle Turonian, but later obser- lateral horns on the adult body chamber. Col- vations by Kennedy in the area around Ci- lignoniceras woollgari is compressed and zay-la-Madelaine, southeast of Saumur in ®nely ribbed in early and middle growth, Touraine, France (see Kennedy and Juignet, with siphonal clavi maintained to maturity. 1981: ®gs. 1, 2) suggest it may extend down Very small specimens (up to 7 mm diameter) into the top of the lower Turonian Mammites of C. woollgari have trituberculate venters nodosoides Zone. Inner whorls of Lecointri- (Cobban and Hook, 1979: pl. 2, ®gs. 18, 19). ceras are characterized by umbilical, feeble At maturity, ribbing coarsens markedly in or no inner ventrolateral, and strong outer both dimorphs of C. woollgari, with large ventrolateral and siphonal tubercles (see umbilical ¯ares, large ventrolateral horns and Kennedy et al., 1980: pl. 74, ®gs. 3±10), fea- a ®nal, nearly smooth, tubular portion of the tures similar to those of Cibolaites. However, adult body chamber, a general style of or- the body chamber characters readily distin- nament similar to adult Lecointriceras, but guish the two genera. In Cibolaites, the ma- with the addition of persistent siphonal clavi. ture body chamber is ¯at-sided with progres- Merewether et al. (1979) drew attention to sively weakening tubercles and ribs. In Le- the occurrence of both early and late forms cointriceras, the ribs and tubercles strength- of C. woollgari in the U.S. Western Interior, en with the development of massive an observation expanded on by Cobban and ventrolateral horns, a ¯attened venter, and tu- Hook (1979), who recognized an earlier C. bular, near-smooth terminal portion immedi- woollgari woollgari and a later C. woollgari ately prior to the adult aperture; adults of the regulare (which are taken as indices of two type species are dimorphic (Kennedy et al., subzones of the woollgari Zone; ®g. 2). They 1984). Lecointriceras thus preserves in its differ in that C. woollgari woollgari has mid- earliest ontogeny features similar to those of dle and late growth stages characterized by Cibolaites, but by its development of inner secondary ribs, more siphonal tubercles than ventrolateral or outer lateral tubercles as well ventrolateral ones, and looped ribs connect-

← Fig. 11. Collignoniceras woollgari woollgari (Mantell, 1822). A, B. USNM 420009, a gracile form from locality 62. C, D. USNM 252793, a gracile form from locality 50. E, F. USNM 252795, a gracile form from locality 52. G, H. USNM 356903, a gracile form from locality 53. I, J. USNM 498215, a gracile form from locality 49. K, L. USNM 420010, a gracile form from locality 62. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_18 Allen Press • DTPro System File # 01235TQ

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Fig. 13. Collignoniceras woollgari woollgari (Mantell, 1822). USNM 498216, from locality 48. Figure is ϫ1.

← Fig. 12. Collignoniceras woollgari woollgari (Mantell, 1822). A. USNM 420006; B, C. USNM 420007, both gracile forms from locality 62. D, E. USNM 403228, a robust form from locality 54. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_20 Allen Press • DTPro System File # 01235TQ

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ing ventrolateral horns, whereas C. woollgari regulare lacks these features in middle growth. Adult body chambers of the two subspecies may, however, be inseparable in some cases. Kennedy et al. (1986) recorded fragments with intercalated ribs from the upper Turonian of Aube, France, but these are bits of body chamber, and restudy of material from Touraine in France shows that the two successive subspecies can also be recognized there, and indeed elsewhere in western Eu- rope. There are several other species of Collig- noniceras that co-occur with Collignoniceras woollgari in western Europe. Collignonicer- as carolinum (D'Orbigny, 1850) (p. 310, pl. 9, ®gs. 5, 6; Kennedy et al., 1980: 574, pl. 68, ®gs. 1±11; pl. 76, ®gs. 1, 2; text-®gs. 1B, 5) is a small species (up to 12 cm adult diameter) that is ®nely and densely ribbed and does not develop hypernodose body chamber ornament; it ranges through much of the Eu- ropean woollgari Zone. Collignoniceras pa- pale (D'Orbigny, 1841) (p. 354, pl. 109, ®gs. 1±3; Kennedy et al., 1980: 578, pl. 69, ®gs. 1, 2; pl. 70, ®gs. 3±5; text-®gs. 1c, 6, 7), is a coarsely ribbed, medium-sized species (up Fig. 14. Collignoniceras woollgari woollgari to 17 cm adult diameter) of the regulare Sub- (Mantell, 1822). USNM 498216, from locality 48. zone that also lacks hypernodose body cham- Figure is ϫ1.

Fig. 15. External suture of Collignoniceras woollgari woollgari (Mantell, 1822), taken from Oxford University Museum of Natural History Collections, no. KT 1275, Ojinaga Formation, middle Turonian, C. woollgari Zone, C. woollgari woollgari Subzone, Cannonball Hill, Chihuahua, Mexico. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_21 Allen Press • DTPro System File # 01235TQ

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Fig. 16. Distribution of Collignoniceras woollgari woollgari (Mantell, 1822), and C. woollgari regulare (Haas, 1946) in the Western Interior Seaway during the early middle Turonian. Land areas indicated by stipple. Modi®ed after Cobban et al., 1994. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_22 Allen Press • DTPro System File # 01235TQ

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Fig. 17. Collignoniceras woollgari regulare (Haas, 1946), gracile form. A±C. USNM 498227; D± F. USNM 498217; G±I. USNM 498228; J, K. USNM 498229; L±N. USNM 498221; O±Q. USNM 498230; R, S. USNM 498231, all from locality 8. T, U. USNM 498232, from locality 12. V, W. USNM 498233, from locality 8. X, Y. USNM 498234, from locality 12. Z±B1. USNM 498223, from locality 8. All ®gures are ϫ1.

ber ornament and is dimorphic (Kennedy et dium-sized species with coarsely ornamented al., 1984). Collignoniceras canthus (Sornay, inner whorls but a weakly ornamented body 1951) (p. 629, text-®gs. 1, 2; Kennedy et al., chamber. Collignoniceras turoniense (Sor- 1980: 582, pl. 73, ®gs. 1±4) is a rare, me- nay, 1951) (p. 630, pl. 21, ®gs. 1±3; Kennedy BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_23 Allen Press • DTPro System File # 01235TQ

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Fig. 18. Collignoniceras woollgari regulare (Haas, 1946), gracile form. A, B. USNM 498235, from locality 12. C, D. USNM 498236; E±G. USNM 498237, both from locality 8. All ®gures are ϫ1.

et al., 1980: 584, pl. 71, ®gs. 4, 5; pl. 72, ing the umbilical ones, and a nearly contin- ®gs. 1±3) is a medium-sized species with uous siphonal keel, whereas in C. woollgari coarse ornament throughout. regulare the number of siphonal clavi equals In the U.S. Western Interior, C. woollgari the number of ventrolateral tubercles except disappears earlier in the middle Turonian in occasional, very large individuals (30 cm than in western Europe, and is replaced by diameter). These features also separate C. four endemic species: Collignoniceras ver- praecox from C. vermilionense, as does the milionense (Meek and Hayden, 1860), C. retention of inner and outer ventrolateral tu- jorgenseni n. sp., C. percarinatum (Hall and bercles in C. praecox at an ontogenetic stage Meek, 1856), and C. praecox (Haas, 1946). when they are absent in C. vermilionense. Collignoniceras vermilionense is a diminu- Collignoniceras jorgenseni is a small species tive, very evolute species, with ribs that lack with prorsiradiate primary and secondary well-de®ned ventrolateral tubercles. It is ribs throughout ontogeny, bullate primary smaller than C. woollgari regulare. Collig- ribs, inner and outer ventrolateral tubercles, noniceras praecox differs from C. woollgari siphonal clavi equal in number to the ribs, regulare by the persistence of long and short and primary ribs that are commonly ¯ared. ribs, with ventrolateral tubercles outnumber- In the middle Turonian Western Interior, BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_24 Allen Press • DTPro System File # 01235TQ

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Fig. 19. Collignoniceras woollgari regulare (Haas, 1946), gracile form. USNM 498238, from locality 17. Figure is ϫ1.

there are two diminutive species that are pre- 6A); it has a phragmocone with umbilical sumed to be paedomorphic in origin. Collig- bullae, outer ventrolateral and siphonal clavi, nonicerites collisniger n. gen., n. sp. is adult and a smoothing, constricted adult body at 18 mm diameter or less and retains the chamber. Collignoniceras percarinatum is very simple suture of juvenile Collignonicer- also a small species; the largest observed as to maturity (Matsumoto, 1965: text-®g. adult is 39 mm in diameter. It most closely BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_25 Allen Press • DTPro System File # 01235TQ

2001 KENNEDY ET AL.: COLLIGNONICERATINAE 25

Fig. 20. Collignoniceras woollgari regulare (Haas, 1946). A. USNM 498238, gracile form from locality 17. B±D. isolated ventrolateral spine, BHI 2103, from the lower part of the Carlile Shale in sec. 12, T 9 S, R 1 E, Fall River County, South Dakota. Collected by N. L. Larson. All ®gures are ϫ1.

resembles the densicostate, evenly ribbed Wright, 1979), occur stratigraphically above variants of woollgari regulare, but retains the C. woollgari regulare, as described by Mat- juvenile ornament of these forms to maturity. sumoto (1965: 16). In western Europe and elsewhere, a series of Whereas Subprionocyclus extends into the super®cially similar species, referred to the uppermost Turonian in Europe, Japan, and genus Subprionocyclus Shimizu, 1932 (see elsewhere, the later Turonian history of the BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_26 Allen Press • DTPro System File # 01235TQ

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Fig. 21. Collignoniceras woollgari regulare (Haas, 1946). A, B. USNM 498239, gracile form. C, BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_27 Allen Press • DTPro System File # 01235TQ

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Collignoniceratinae in the Western Interior is gascar, Venezuela, and Colombia, and pos- dominated by a succession of species of sibly from Texas; and Gauthiericeras de Prionocyclus Meek, 1876, most of which are Grossouvre, 1894, of the upper Coniacian, is endemic to the region or are known only as known as a rarity in the Western Interior in rare individuals in North Africa, western Eu- Montana and New Mexico (Kennedy and rope, and Japan. These are medium-sized to Cobban, 1991), as well as occurring in Eu- large taxa, with adults up to 300 mm in di- rope, north, east, and west Africa, Madagas- ameter. We have failed to recognize size- car, New Caledonia, Mexico, Colombia, linked dimorphism in this genus, due to pau- Peru, Venezuela, and Argentina (Wright, city of complete adults. Ornament is highly 1996). variable. The type species of Collignoniceras and Prionocyclus are very distinct (see be- VARIATION WITHIN THE low), and juveniles can be distinguished because the keel of Collignoniceras has si- COLLIGNONICERATINAE phonal clavi equal in number to the ventro- Haas (1946) observed considerable intra- lateral tubercles, whereas in Prionocyclus the speci®c variation within what he referred to serrations outnumber the ribs. Prionocyclus as Prionotropis woollgari Meek (? non Man- also differs from Collignoniceras in the per- tell). This form of the species, later referred sistence to maturity of the marked differen- to as Collignoniceras woollgari regulare tiation of ribbing of the type shown by cer- Haas, 1946, by Cobban and Hook (1979) and tain variants of C. praecox when young. Fi- Merewether et al. (1979), was subdivided by nally, Prionocyclus is characterized by the Haas into the forma typica, variety crassa, appearance of the keel in early ontogeny. var. intermedius, var. regularis, var. tenuicos- In addition to Prionocyclus hyatti, there is tata, var. praecox, and var. alata. These var- a succession of species that differ in details iants were based on details of the ontogeny, of ornament, as described in the systematic ornamentation, and sutures. Haas noted gra- section below: P. albinus (Fritsch, 1872), P. macombi Meek, 1876, P. bosquensis Ken- dation between most of the named forms ex- nedy, 1988, P. wyomingensis Meek, 1876, P. cept his variety C. woollgari praecox, which novimexicanus (Marcou, 1858), P. quadratus he thought could be a separate species. Most Cobban, 1953, P. germari (Reuss, 1845), and of Haas's material came from a single lime- P. pluricostatus n. sp. Prionocyclites mite stone concretion from the lower part of the Kennedy, 1988, from the P. hyatti Zone of Mancos Shale of southern Utah. Texas is a diminutive species, reminiscent of During the ®rst of his visits to the United Collignonicerites collisniger n. gen., n. sp., States, Tatsuro Matsumoto was taken by one of the preceding C. woollgari regulare Sub- of us (WAC) on a tour of the Cretaceous zone. rocks of the Black Hills area in eastern Wy- Reesidites Wright and Matsumoto, 1954, is oming and western South Dakota. Matsu- rare and only known from a few specimens moto made a large collection of C. woollgari in the Prionocyclus wyomingensis and Sca- (now assigned to C. woollgari regulare) phites whit®eldi zones of New Mexico (Cob- from limestone concretions in the lower part ban and Kennedy 1988). There are no early of the Carlile Shale at USGS Mesozoic lo- Coniacian Collignoniceratinae known from cality 21792 near Newcastle, Wyoming. the Western Interior. Prionocycloceras Spath, Matsumoto was impressed by the great var- 1926, of the Coniacian is known with cer- iation within the species, and in 1965 Mat- tainty from Armenia, North Africa, Mada- sumoto assigned his specimens to six groups

← D. USNM 498220, robust form. E±G. USNM 498218, gracile form. H, I. USNM 498219, gracile form. J±L. USNM 498240, robust form. M±O. USNM 498241, gracile form. P±R. USNM 498222, gracile form. S, T. USNM 498242, robust form. U±W. USNM 498224, gracile form. X±Z. USNM 498243, gracile form. All specimens are from locality 8. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_28 Allen Press • DTPro System File # 01235TQ

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lettered A±F, based largely on variation in mensions are given in millimeters, with D ϭ ornamentation. diameter; Wb ϭ whorl breadth; Wh ϭ whorl Regarding the genus Prionocyclus, Haas height; and U ϭ umbilical diameter (®g. 4). (1946) noted intraspeci®c variation within P. In the tables, values in parentheses refer to wyomingensis Meek, 1876. Haas grouped the dimensions as a percentage of diameter. variations as the forma typica, the variety Specimens are photographed in the custom- elegans, and the variety robusta. His variety ary position with the aperture on top al- elegans was later determined to be a syno- though the authors recognize that the animal nym of Ammonites novimexicanus Marcou, would have been oriented differently in life. 1858, by Hook and Cobban (1979), who Arrows on photographs indicate the adapical demonstrated that Marcou's form was a end of the body chamber, if preserved. younger species than P. wyomingensis. The present authors have observed consid- SYSTEMATIC PALEONTOLOGY erable intraspeci®c variation in all of the FAMILY COLLIGNONICERATIDAE WRIGHT AND larger collections of Collignoniceratinae WRIGHT, 1951 studied. Most species seem to consist of two major forms, a gracile one that has whorls higher than wide, and a more robust form [pro Prionotropidae Zittel, 1895: 430, ex that has broader whorls, a wider umbilicus, Prionotropis Meek, 1876a: 453, non Fieber, and stronger ornamentation. These may be 1853: 127; ϭ Prionocyclidae Breistroffer, sexual dimorphs, but we generally lack suf- 1947, unpaged, ex Prionocyclus Meek, 1871: ®cient complete adult individuals to deter- 298, ineligible as family type] mine if robust and gracile variants differ in adult size. In the following systematic treat- SUBFAMILY COLLIGNONICERATINAE ments, these are referred to as gracile and WRIGHT AND WRIGHT, 1951 robust forms. Genus Cibolaites Cobban and Hook, 1983

CONVENTIONS TYPE SPECIES: Cibolaites molenaari Cob- ban and Hook, 1983: 16, pl. 2, ®gs. 1±9; pl. The following abbreviations are used to 3, ®gs. 3±8; pl. 8, ®gs. 6±8; pl. 13, ®gs. 1± indicate the repositories of specimens cited 5; pl. 14; text-®gs. 13, 14; lower Turonian in the text: Mammites nodosoides Zone, west-central AMNH American Museum of Natural History, New Mexico. New York DIAGNOSIS: A medium-size genus, largest BMNH Natural History Museum, London known adults 120 mm in diameter; dimor- OUM Oxford University Museum of Natural phism not demonstrated. Coiling involute History, Oxford with small umbilicus, whorl section com- TMM Texas Memorial Museum, Austin pressed to depressed, intercostal section BHI Black Hills Museum of Natural Histo- with convergent sides and fastigiate venter, ry, Hill City, South Dakota USNM U.S. National Museum of Natural His- costal section markedly concave between tory, Washington, D.C. tubercles. Strong conical umbilical bullae give rise to primary ribs singly or in pairs The system of sutural terminology of We- with shorter, intercalated ribs, all with ven- dekind (1916) as propounded by Kullmann trolateral and siphonal clavi on phragmo- and Wiedmann (1970) is used here, with E cone. Umbilical bullae decline on the adult ϭ external lobe; L ϭ lateral lobe; U ϭ um- body chamber, which is ¯at-sided with bilical lobe; and I ϭ internal lobe. All di- crowded weak primary and intercalated

← Fig. 22. Collignoniceras woollgari regulare (Haas, 1946). A±C. USNM 498244, robust form from locality 8. D, E. USNM 498245; F. USNM 498246; G, H. USNM 498247, all robust forms from locality 7. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_30 Allen Press • DTPro System File # 01235TQ

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Fig. 23. Collignoniceras woollgari regulare (Haas, 1946). A±C. USNM 498226, gracile form from locality 8. D, E. USNM 252723, gracile form from locality 7. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_31 Allen Press • DTPro System File # 01235TQ

2001 KENNEDY ET AL.: COLLIGNONICERATINAE 31

Fig. 24. Collignoniceras woollgari regulare (Haas, 1946). USNM 420012, robust form from locality 62. Figures are ϫ0.80.

ribs; ventrolateral and siphonal tubercles noniceratinae. Cibolaites super®cially resem- decline progressively and the venter ¯at- bles Barroisiceras de Grossouvre, 1894, and tens towards the adult aperture. Subbarroisiceras Basse, 1946, of the upper Suture relatively simple with broad, little- Turonian±Coniacian Barroisiceratinae. How- divided saddles and narrower, rectangular ever, they differ in sutural characteristics, lobes. while both are more involute than Cibolaites, DISCUSSION: Coarse tuberculation and with smaller umbilici and crenulate keels; in presence of only a single row of ventrolateral Cibolaites there is no continuous keel and the tubercles distinguishes Cibolaites from all siphonal clavi are separated by smooth inter- other genera currently referred to the Collig- spaces. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_32 Allen Press • DTPro System File # 01235TQ

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Fig. 25. Collignoniceras woollgari regulare (Haas, 1946). USNM 498248, robust form from locality 14. Figures are ϫ0.80. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_33 Allen Press • DTPro System File # 01235TQ

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Fig. 26. Collignoniceras woollgari regulare (Haas, 1946). A, B. MuseÂum National d'Histoire Na- turelle, Paris, no. 6778 (ex d'Orbigny Collection), from the Middle Turonian of PonceÂ, Sarthe, France. C. FaculteÂ des Sciences, Rennes, no. C273, also from PonceÂ. D, E. Same collection as A, B, no. W1, from ``Le Mans, Sarthe,'' France. All ®gures are ϫ1.

OCCURRENCE: Upper Cenomanian Neocar- middle Turonian in west-central New Mexi- dioceras juddii Zone in Devon, England and co; M. nodosoides Zone, N.W. Germany; Aube, France; lower Turonian Mammites no- also recorded from northern Spain (Kaplan, dosoides Zone, immediately below ®rst ap- 1988) and from Tunisia (Chancellor et al., pearance of C. woollgari woollgari of the 1994). BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_34 Allen Press • DTPro System File # 01235TQ

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Fig. 27. Collignoniceras woollgari regulare (Haas, 1946). A, B. BMNH 4863, from ``near Lewes,'' Sussex, England (ex J. de C. Sowerby Collection). C, D. BMNH 4863a, from the same locality. E. paralectotype BMNH 5742b. F. Paralectotype BMNH 5742a, both from near Lewes, Sussex. G. BMNH C82268, from the Middle Turonian Middle Chalk, upper Terebratulina lata Zone, Mickleham Bypass, Surrey, England. All ®gures are ϫ1.

Cibolaites molenaari Cobban and Hook, Cibolaites sp. Kaplan, 1988: 23, pl. 6, ®gs. 4, 5. 1983 Cibolaites molenaari Cobban and Hook. Cobban Figures 5±8 and Hook, 1989: ®gs. 8c, d. Cibolaites molenaari Cobban and Hook, 1983. Collignoniceras sp. Wright and Kennedy, 1981: Reyment and Kennedy, 2001: ®g. 1a, b. 107, pl. 8, ®g. 17. Cibolaites molenaari Cobban and Hook, 1983: TYPES: Holotype is USNM 328766 (®g. 5), 16, pl. 2, ®gs. 1±9; pl. 3, ®gs. 3±8; pl. 8, ®gs. from the Rio Salado Tongue of the Mancos 6±8; pl. 13, ®gs. 1±5; pl. 14; text-®gs. 13, 14. Shale, lower Turonian Mammites nodosoides Cibolaites molenaari Cobban and Hook. Kennedy Zone, immediately below the ®rst occurrence et al., 1986: 209, text-®g. 5i, j, k, l. of Collignoniceras woollgari woollgari of

→ Fig. 28. Collignoniceras woollgari regulare (Haas, 1946). MuseÂum National d'Histoire Naturelle, Paris, no. W2. 1904±32, from ``Le Mans, Sarthe,'' France. Figures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_35 Allen Press • DTPro System File # 01235TQ

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Fig. 29. Collignoniceras woollgari regulare (Haas, 1946). MuseÂum National d'Histoire Naturelle, Paris, 102±27, from the Middle Turonian of PonceÂ, Sarthe, France. Reduced ϫ0.625.

the middle Turonian at USGS Mesozoic lo- moderate-size species are the strong, nodate cality D11208, NE1/4 sec. 36, T. 6 N, R. 19 umbilical tubercles during early and middle W, Cibola County, New Mexico. Paratypes growth and the equal numbers of prominent USNM 328752±328767, 329013, are from siphonal clavi and clavate inner ventrolateral the Rio Salado Tongue at this and nearby lo- tubercles. calities. DESCRIPTION: The early whorls are smooth DIAGNOSIS: Characteristic features of this to a diameter of 5 mm, with rounded ¯anks BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_37 Allen Press • DTPro System File # 01235TQ

2001 KENNEDY ET AL.: COLLIGNONICERATINAE 37

Fig. 30. Collignoniceras woollgari regulare (Haas, 1946). MuseÂum National d'Histoire Naturelle, Paris, W. 22, 6778 (ex d'Orbigny Collection), from the Middle Turonian of PonceÂ, Sarthe, France. Reduced ϫ0.625. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_38 Allen Press • DTPro System File # 01235TQ

38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 267

Fig. 31. Collignoniceras woollgari regulare (Haas, 1946). MuseÂum National d'Histoire Naturelle, Paris, 1946±19, from the Middle Turonian of Sainte-Maure-de-Touraine, Indre et Loire, France. Reduced approximately ϫ0.7.

and venter. Low umbilical bullae, 14 per (®gs. 6, 7), the number of bullae decreases whorl, appear between 5 and 7 mm diameter; to as few as seven per whorl in middle ventrolateral and siphonal tubercles appear growth. Coiling is fairly involute, with a around 10 mm diameter. As size increases deep umbilicus. Two broad prorsiradiate to BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_39 Allen Press • DTPro System File # 01235TQ

2001 KENNEDY ET AL.: COLLIGNONICERATINAE 39

Fig. 32. Collignoniceras woollgari regulare (Haas, 1946). MuseÂum National d'Histoire Naturelle, Paris, 1946±19, from the Middle Turonian of Sainte-Maure-de-Touraine, Indre et Loire, France. Reduced approximately ϫ0.7.

rectiradiate ribs generally arise from each vi; occasional intercalated ribs arise around bulla, which vary in strength within and be- mid¯ank or below and bear clavi of compa- tween specimens. The ribs are broad, round- rable strength to give a total of 22 or so ribs ed, and terminate in strong ventrolateral cla- per whorl at diameters of up to 65 mm. Each BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_40 Allen Press • DTPro System File # 01235TQ

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ical bullae decline and ultimately disappear. The ventrolateral and siphonal clavi weaken markedly, but are still present, though very weak, immediately adapical of the adult aperture. As they decline, the venter broadens and rounds. On the body chamber, the ¯anks ¯atten and the whorl section becomes rect- Fig. 33. External suture of Collignoniceras angular, with low, weak, prorsiradiate ribs, woollgari regulare (Haas, 1946). Copy of Haas, irregularly long and short, the latter of vari- 1946: ®gure 83. able length. Part or all of the body chamber is preserved on 46 specimens. Diameter at the last septum ranges from 44 to 94 mm, ventrolateral clavus gives rise to a broad, low without any clear clustering into size classes prorsiradiate rib that arches forward across or indication of size dimorphism. Suture with the venter to form an obtuse chevron with a broad, bi®d E/L, narrow, bi®d L, slightly strong siphonal clavus at the apex. In pro®le, broader L/U2, and much narrower U2 (®g. 8). the adapical slope of this clavus is low, the Cibolaites molenaari occurs as gracile and adapertural slope much steeper, giving the robust forms, as indicated in the ®gure cap- clavus a distinctly asymmetric pro®le. At this tions, although the distinctions between them stage the costal and intercostal whorl sections are not as pronounced as in species of Col- vary from compressed to depressed (see table lignoniceras and Prionocyclus. The holotype 1); the greatest breadth in intercostal section and paratypes ®gured by Cobban and Hook is just outside the umbilical shoulder. The in- (1983) can be assigned to the following ner ¯anks are broadly rounded, the outer forms: gracile form, pl. 2, ®gs. 1±6, pl. 3, ¯anks ¯attened and convergent, and the ven- ®gs. 3, 4, 6±8, pl. 14, ®gs. 1±15; robust form, ter fastigiate. The costal section has the pl. 2, ®gs. 7±9, pl. 3, ®g. 5, pl. 8, ®gs. 6±8, greatest breadth at the umbilical bullae and pl. 13, ®gs. 1±5. the ¯anks are concave between tubercles; the DISCUSSION: Coarse ornament and pres- venter is markedly fastigiate. On the later ence of only a single row of ventrolateral tu- parts of the phragmocone as well as on the bercles at all stages distinguishes C. mole- adult body chamber, the ornament is pro- naari from all of the collignoniceratids de- gressively modi®ed (®g. 5); ®rst, the umbil- scribed here. Body chamber characters sep-

TABLE 2 Dimensions of Collignoniceras woollgari regulare (Haas, 1946)a BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_41 Allen Press • DTPro System File # 01235TQ

2001 KENNEDY ET AL.: COLLIGNONICERATINAE 41

Fig. 34. A±C. Collignoniceras jorgenseni n. sp., USNM 498250, a paratype, robust form from locality 20. D±H. Collignoniceras vermilionense (Meek and Hayden, 1860). D. cast of holotype, USNM 224; E. Copy of original ®gure of Meek, 1876a: plate 7, ®gure 2. Original is from a limestone concretion in the Carlile Shale, probably the middle Turonian Collignoniceras praecox Zone, on the Missouri River opposite the mouth of Vermillion (formerly Vermilion) River in Dixon County, Nebraska. F, G. USNM 299166, gracile form from locality 21. H. S. Jorgensen Collection CO1970a, Carlile Shale, Collignon- iceras praecox Zone, dredged from Missouri River during natural gas pipeline construction at Yankton, South Dakota. All ®gures are ϫ1.

arate it from Lecointriceras ¯euriausianum ren and Stelck, 1940: 151; non Collignoni- (D'Orbigny, 1841) (®g. 9) as discussed ceras Van Hoepen, 1955: 361]. above. TYPE SPECIES: Ammonites woollgari Man- OCCURRENCE: Upper Cenomanian Neocar- tell, 1822: 197, pl. 21, ®g. 16; pl. 22, ®g. 7, dioceras juddii Zone in Devon, England, and by the original designation of Meek, 1876a: Aube, France; lower Turonian Mammites no- 453, as type species of Prionotropis Meek, dosoides Zone in west-central New Mexico 1876a (non Fieber, 1853) for which Breis- and northeastern Arizona; N.W. Germany. troffer (1947, unpaged) proposed Collignon- iceras as nomen novum, from the Turonian Genus Collignoniceras Breistroffer, 1947 of England. DIAGNOSIS: ``Medium-sized, moderately [ICZN, 1968, Opinion 861, name no. involute to evolute ammonites. Early whorls 1798; pro Prionotropis Meek, 1876a: 453, compressed, parallel-sided, ornamented by non Fieber, 1853: 127; ϭ Selwynoceras War- crowded or sparse, prorsiradiate, straight or BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_42 Allen Press • DTPro System File # 01235TQ

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or ¯ared rib, from which commonly arise pairs of low ribs joining siphonal clavi more numerous than the ventrolateral and linked into a more or less continuous keel. Rarely the ornament is greatly reduced on the body whorl. Sutures little incised, with massive saddles'' (Wright and Kennedy, 1981: 102). DISCUSSION: See Kennedy et al. (1980: 558) and Wright and Kennedy (1981: 102). OCCURRENCE: Middle to lower upper Tu- ronian, England, France, Germany, Czech Republic, Poland, Roumania, eastwards to Turkmenistan, Japan, California, northern Mexico, the U.S. and Canadian Interiors, Greenland, North Africa, Colombia, south India, northern Australia.

Collignoniceras woollgari woollgari (Mantell, 1822) Fig. 35. A. Partial external suture of Collig- Figures 10±15 noniceras vermilionense (Meek and Hayden, 1860). S. Jorgensen Collection no. CO1970A Ammonites woollgari Mantell, 1822: 197, pl. 21, from the Middle Turonian C. praecox Zone, con- ®g. 16; pl. 22, ®g. 7. cretion dredged from Missouri River during nat- Collignoniceras woolgari (sic) (Mantell) (early ural gas pipeline construction at Yankton, South form). Merewether et al., 1979: pl. 3, ®gs. 9, Dakota. B. Collignoniceras jorgenseni, n. sp. Par- 10. tial external suture of paratype USNM 498253, Collignoniceras woollgari woollgari (Mantell). collected by S. D. Jorgensen from the Middle Tu- Cobban and Hook, 1979: 21, pl. 1, ®gs. 1±11; ronian C. praecox Zone, Carlile Shale at locality pl. 2, ®gs. 5±22; pl. 4, ®gs. 11, 12; pl. 5, ®gs. 16. 13±16; pl. 12, ®gs. 1, 2 (with synonymy). Collignoniceras woollgari (Mantell). Kennedy et al., 1980: 560 (pars), pl. 62; pl. 63, ®gs. 1±4, ¯exuous ribs, mostly long, with weak to 7±12, non 5, 6; pl. 64; pl. 65, ®gs. 1±3; text- strong umbilical bullae. All ribs bear in the ®g. 2, ?3, 4c, d, non 4a, b. early stages outer ventrolateral tubercles in Collignoniceras woollgari (Mantell, 1822). addition to siphonal clavi. This style of or- Wright and Kennedy, 1981: 103, pl. 28, ®g. 3, non 1, 2; pl. 29, ®g. 5, non 1±4, 6, 7; pl. 30, nament is, in some species, retained to ma- ®g. 1, non 2, 3. turity. In most, however, the ribs coarsen, be- Collignoniceras woollgari (Mantell, 1822). come widely spaced, with strong to weak AmeÂdro and Badillet, 1982: 131, pl. 5, ®g. 1. umbilical tubercles (which migrate progres- Collignoniceras woollgari (Mantell). AmeÂdro et sively outward from the umbilical margin), al., 1982: 30, pl. 1, ®gs. 5±7. prominent inner and outer ventrolateral tu- Collignoniceras woollgari (Mantell). AmeÂdro and bercles, which may fuse into a massive horn Hancock, 1985: ®g. 9e, f.

TABLE 3 Dimensions of Collignoniceras vermilionense (Meek and Hayden, 1860)a BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_43 Allen Press • DTPro System File # 01235TQ

2001 KENNEDY ET AL.: COLLIGNONICERATINAE 43

Fig. 36. Collignoniceras jorgenseni, n. sp. A. holotype, USNM 498249, robust form; B±D, K. Paratype USNM 498253, gracile form; both from locality 16. E±G. USNM 356918, robust form from the lower part of the Carlile Shale, center of sec. 35, T 8S, R 1E, Fall River County, South Dakota. H± J. Paratype USNM 498254, gracile form from locality 16. All ®gures are ϫ1.

Collignoniceras woollgari woollgari (Mantell). lan, 1988: 11, pl. 1, ®g. 1; pl. 2, ®gs. 1±7; pl. Cobban, 1986: 81, ®gs. 6j, k; 7k, l. 7, ®g. 1. Collignoniceras woollgari woollgari (Mantell, Collignoniceras woollgari woollgari (Mantell, 1822). Kennedy and Cobban, 1988: 604, ®gs. 1822). Kennedy et al., 1989: 86, ®g. 24a±c, h±l. 7.14, 15. Collignoniceras woollgari (Mantell, 1822). Cob- Collignoniceras woollgari (Mantell, 1822). Kap- ban and Hook, 1989: ®g. 9q, r. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_44 Allen Press • DTPro System File # 01235TQ

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2001 KENNEDY ET AL.: COLLIGNONICERATINAE 45

Collignoniceras woollgari (Mantell, 1822). Collignoniceras woollgari regulare (Haas, AmeÂdro, 1990: 269, pl. 24, ®g. 2. 1946) Collignoniceras woollgari woollgari (Mantell, Figures 17±33, 49L 1822). Emerson et al., 1994: 208, unnumbered ®gure. Prionocyclus (Prionotropis) woolgari (sic) (Man- Collignoniceras woollgari woollgari (Mantell, tell). Meek, 1876a: 455, pl. 6, ®g. 2; pl. 7, ®gs. 1822). Reyment and Kennedy, 2001: ®g. 1c, d. 1, 3. Prionocyclus (Prionotropis) woolgari (sic) (Man- tell). Boyle, 1893: 243. TYPES: Lectotype (®g. 10) is BMNH 5682, Prionotropis woolgari (sic) Mantell. Stanton, the original of Mantell, 1822: pl. 21, ®g. 16, 1894: 174, pl. 42, ®gs. 1±4. from the Middle Chalk of Lewes, Sussex, Prionotropis woolgari (sic) Mantell. Logan, 1898: England; designated by Wright and Wright, 466, pl. 102, ®gs. 1±4. Prionotropis woolgari (sic) Mantell. Roman, 1951: 35. Presumed paralectotypes (®g. 27E, 1938: 455, pl. 46, ®g. 434b, c. F) are BMNH C5742a, b, from the same lo- Prionotropis graysonensis (Shumard). Moreman, cality. 1942: 213 (pars). DIAGNOSIS: This moderately evolute sub- Prionotropis woolgari (sic) (Mantell). Shimer and species has prorsiradiate to rectiradiate pri- Shrock, 1944: 593, pl. 247, ®gs. 1, 2. mary and secondary ribs that support bullate Prionotropis woollgari (Mantell) and varieties. inner ventrolateral tubercles and clavate out- Haas, 1946: 150, pls. 11, 12; pl. 13, ®gs. 1±3, 5±18 (non 4, 19); pl. 14, ®gs. 1±10, 12±16 (non er ventrolateral tubercles matched by siphon- 11); pl. 15, ®gs. 1±10; pl. 16, ®gs. 1±21 (non al clavi. 22±33); non pl. 17; pl. 18, ®gs. 1, 3±7 (non 8, DISCUSSION: Cobban and Hook (1979) pro- 9); text-®gs. 1±4, 6±14, 19±79, 80±83, 91. vided an account of C. woollgari woollgari Prionotropis Adkins and Lozo, 1951: pl. 5, ®gs. from New Mexico, and Kennedy et al. 4±9. (1980) and Wright and Kennedy (1981) re- Collignoniceras woollgari (Mantell). Cobban et viewed the European material at length. We al., 1956: 1270, ®g. 1b±h. illustrate here a selection of European, Mex- Collignoniceras woollgari (Mantell). Matsumoto and Miller, 1958: 353, pl. 44, ®gs. 1±6; pl. 45, ican, and U.S. Western Interior material (®gs. ®g. 1. 10±15) for comparison with C. woollgari re- Collignoniceras woollgari (Mantell). Matsumoto, gulare, described below, where features dis- 1959: 105, ®gs. 55±57. tinguishing it from the nominate subspecies Collignoniceras woollgari (Mantell). Hattin, are outlined. Gracile and robust forms of C. 1962: pl. 12, ®gs. D, E. woollgari woollgari are present, as indicated Collignoniceras woollgari (Mantell). Hattin, in the ®gure captions. Specimens ®gured by 1965: ®g. 4±3. Collignoniceras woollgari (Mantell). Matsumoto, Cobban and Hook (1979) include gracile 1965: 11, pl. 1, ®gs. 1±6; pl. 2, ®gs. 1±3; pl. 3, forms (pl. 1, ®gs. 1±3, 10, 11; pl. 2, ®gs. 5± ®gs. 1, 2; text-®g. 6. 19; pl. 5, ®gs. 13±16), and robust forms (pl. Prionocyclus (Collignoniceras) woolgari (sic) 1, ®gs. 4±9; pl. 2, ®gs. 20±22; pl. 4, ®gs. 11, (Mantell). Jeletzky, 1970: pl. 26, ®g. 9a, b. 12; pl. 12, ®gs. 1, 2). Prionocyclus (Prionocyclus) wyomingensis Meek OCCURRENCE: Collignoniceras woollgari var. robusta Haas. Jeletzky, 1970: pl. 26, ®g. 3. woollgari Subzone, widespread in the U.S. Prionocyclus (Prionocyclus) wyomingensis Meek var. elegans Haas. Jeletzky, 1970: pl. 6, ®g. 8a, Western Interior from Wyoming to the Rio b. Grande; California, northern Mexico, Eu- Collignoniceras woollgari (Mantell). Cobban and rope, east to Turkmenistan; Tunisia, Japan, Scott, 1972: 94, pl. 14, ®g. 5; pl. 30, ®g. 1; pl. and northern Australia. 37, ®gs. 9, 10.

← Fig. 37. Collignoniceras jorgenseni, n. sp. (A) and Collignoniceras vermilionense (Meek and Hay- den, 1860) (B). USNM 498255, from locality 16. Figure is ϫ0.95. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_46 Allen Press • DTPro System File # 01235TQ

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2001 KENNEDY ET AL.: COLLIGNONICERATINAE 47

TABLE 4 Dimensions of Collignoniceras jorgenseni n. sp.a

Collignoniceras woollgari (Mantell). Hattin, Collignoniceras woollgari regulare (Haas). Ken- 1975a: pl. 10, ®gs. N, P±R. nedy and Cobban, 1988: 606, ®gs. 7.9±7.15. Collignoniceras woollgari (Mantell). Hattin, Collignoniceras woollgari (Mantell, 1822) regu- 1975b: pl. 2, ®gs. 1±3. lare (Haas, 1946). Kennedy et al., 1989: 88, Collignoniceras woollgari (Mantell). Cobban, ®gs. 21, 22, 23, 24d. 1976: 120, pl. 1, ®g. 7. Collignoniceras woollgari (Mantell). Kennedy, Collignoniceras woollgari (Mantell). Kennedy 1989: 257, ®g. 7. and Cobban, 1976: 46, pl. 10, ®gs. 1, 2. Collignoniceras woollgari (Mantell). Cobban, Collignoniceras woollgari (Mantell). Kauffman, 1990: B9, pl. 5, ®gs. 6±13. 1977: pl. 22, ®gs. 4±6, pl. 26, ®g. 6. Collignoniceras woollgari regulare (Haas, 1946). Collignoniceras woollgari (Mantell). Hattin, Emerson et al., 1994: 209, 378. 1977: 189, ®g. 8.5. Collignoniceras woollgari regulare (Haas, 1946). Collignoniceras woollgari Mantell. Kauffman et Reyment and Kennedy, 2001: ®g. 1e±j. al., 1978: pl. 5, ®gs. 4±6. Collignoniceras woollgari (Mantell). Hattin and TYPES: Holotype, by original designation, Siemers, 1978: ®gs. 7.10, 10.5. South Dakota State School of Mines Collec- Collignoniceras woollgari (Mantell) (late form). tion no. 1470, the original of Haas, 1946, Merewether et al., 1979: pl. 3, ®gs. 4, 5. plate 16, ®gures 14, 16; text-®gs. 80, 81, Collignoniceras woollgari regulare (Haas). Cob- from an unknown horizon and locality in the ban and Hook, 1979: 22, pl. 3, ®gs. 1±14; pl. Black Hills area (®de Haas, 1946: 198). 12, ®g. 3. MATERIAL: The present description is Collignoniceras woollgari (Mantell). Kennedy et al., 1980: 560 (pars), pl. 63, ®gs. 5, 6; pl. 65, based on more than 500 specimens from ®gs. 4±8. USGS Mesozoic locality 21792, limestone Collignoniceras woollgari (Mantell, 1822). concretions 18.9±21.3 m above the base of Wright and Kennedy, 1981: 103 (pars), pl. 28, the Carlile Shale, middle Turonian Collig- ®g. 2; pl. 29, ®gs. 2, 6, 7; pl. 30, ®g. 3. noniceras woollgari Zone, regulare Sub- Collignoniceras woollgari regulare (Haas). Cob- zone, 1.6 km west of Newcastle in the NW¼ ban, 1983: 16, pl. 15, ®gs. 2±4, 7±11. sec. 31, T. 45 N, R. 61 W, Weston County, Collignoniceras woollgari regulare (Haas). Cob- Wyoming, and from USGS Mesozoic local- ban, 1984b: 14, pl. 2, ®gs. 6, 7. ity D9896, limestone concretions about 18.3 Collignoniceras woollgari (Mantell). Kennedy, 1986: pl. 13, ®gs. 1±3. m below the Turner Sandy Member of the Collignoniceras woollgari (Mantell). Wright and Carlile Shale, middle Turonian Collignoni- Kennedy, 1987: 166, pl. 34, ®gs. 5, 6. ceras woollgari Zone, regulare Subzone, Collignoniceras woollgari regulare (Haas). Ken- NW¼ sec. 35, T. 46 N, R. 63 W, Weston nedy, 1988: 74, pl. 14, ®gs. 1, 2, 4, 5, 10, 11. County, Wyoming. We have several hundred

← Fig. 38. Collignoniceras jorgenseni n. sp. BHI 2102, mass occurrence slab, with associated Sca- phites patulus and Inoceramus sp. Carlile Shale, C. praecox Zone, center of sec. 35, T 8S, R 1E, Fall River County, South Dakota. Figure is ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_48 Allen Press • DTPro System File # 01235TQ

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Fig. 39. Collignoniceras praecox (Haas, 1946), gracile form. A±C. USNM 498268; D±F. USNM 498260; G±I. USNM 498262; J, K. USNM 498259; L, M. USNM 498269; N, O. USNM 498270; P, BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_49 Allen Press • DTPro System File # 01235TQ

2001 KENNEDY ET AL.: COLLIGNONICERATINAE 49

additional specimens from many localities in clavate outer ventrolateral tubercles from an South Dakota, Wyoming, Colorado, Utah, early stage. The siphonal keel is coarse. and New Mexico. There is every intermediate between these DIAGNOSIS: A moderately evolute subspe- two forms in terms of rib density, and great cies that usually has prorsiradiate single ribs variation in the diameter to which the simply that support umbilical and inner and outer ornamented stage persists. Thus, at a diam- ventrolateral tubercles matched by prominent eter of as little as 7 mm (®g. 21M±O) to a siphonal clavi. diameter of up to 27 mm there is a marked DESCRIPTION: Nuclei are highly variable change in rib density; compressed gracile in- (®gs. 17, 21). At one extreme are com- dividuals may have as few as 17 or 18 ribs pressed, high-whorled individuals of the per whorl at diameters of 20±35 mm (®g. gracile form (®g. 17A±C, J, K, O±Q, T, U, 21X±Z), decreasing to as few as 11 by a di- X, Y) with costal whorl breadth to height ameter of 75 mm (®g. 21U±W). The coiling ratios of as little as 0.51 and an umbilical in these gracile specimens is very evolute, to shell diameter ratio of 0.29 (table 2). The with U/D up to 0.38 and intercostal whorl umbilicus is broad, with a low umbilical breadth to height ratios of as little as 0.57 wall, and a broadly rounded umbilical (table 2). The umbilicus is very shallow, with shoulder. The ¯anks are ¯attened and sub- a low umbilical wall, and rounded umbilical parallel with the ventrolateral shoulders shoulder. The ¯anks are ¯attened and sub- rounded. At diameters up to a maximum of parallel in intercostal section with rounded 27 mm, there are dense, crowded ribs that ventrolateral shoulders, and a high, sharp total as many as 70 per whorl. They arise keel. The ribs are very narrow, and the in- singly or in pairs at the umbilical shoulder terspaces between them become progressive- or intercalate, arising at various levels just ly wider as the diameter increases. At small outside the umbilical shoulder and well be- diameters, the ribs originate at umbilical bul- low mid¯ank. The ribs are straight, narrow lae, but with increasing diameter, the ribs be- and rounded, and prorsiradiate to the ven- come very high and ¯ared, reaching their trolateral shoulder, where they ¯ex for- maximum elevation across the ¯anks. The ward; they are concave over the shoulder ribs are markedly prorsiradiate, straight, and and project forward to the siphonal keel, feebly concave across the ¯anks, and each producing an acute chevron. Small outer rib bears a strong conical inner ventrolateral ventrolateral clavi are present from the ear- tubercle and a strong clavate outer ventrolat- liest ornamented stage, and are separated eral tubercle. The two tubercles progressive- by a ventral groove from the siphonal keel, ly assimilate into a ¯ared ventrolateral rib which bears small clavi, equal in number to that sweeps forward over the ventrolateral but displaced slightly aperturally of the shoulder before weakening abruptly prior to outer ventrolateral clavi. Weaker inner ven- meeting the siphonal keel. The keel is high trolateral clavi appear after the outer ones, and crenulate, with long, high, siphonal clavi and may be absent to a diameter of as much that are markedly asymmetric in pro®le (®g. as 12 mm. 23A±E). Fine lirae are developed, or not, on At the other extreme are more robust nu- the interspaces between ribs (®g. 23), but do clei (®g. 21C, D, J±L) with whorl breadth to not bear any tubercles. There are an equal height ratios of up to 0.95 and an umbilical number of umbilical bullae, inner and outer to shell diameter ratio of up to 0.40 (table 2). ventrolateral, and siphonal tubercles. The These have as few as 24 ribs per whorl, all largest individual of the gracile type from the of which are coarse, blunt, straight and pror- Carlile Shale concretions has a diameter of siradiate. The ribs bear weak umbilical bullae 105 mm; fragments have whorl heights of up and prominent conical inner ventrolateral and to 150 mm.

← Q. USNM 498264; R, S. USNM 498261; T. USNM 498271; U±W. USNM 498265; all from locality 15. X±Z. USNM 498272 from locality 12. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:35 PM 2000 amnb 01235 Mp_50 Allen Press • DTPro System File # 01235TQ

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2001 KENNEDY ET AL.: COLLIGNONICERATINAE 51

Robust variants show a comparable ae. We tentatively interpret these forms as change in ribbing density at variable diame- adult microconchs. ters. At diameters of up to 75 mm, these Much larger body chambers (®gs. 24, 25) specimens have umbilical to shell diameter belong to the robust form. A South Dakota ratios of up to 0.34 and a quadrate whorl example is shown in ®g. 25; at Chispa Sum- section. Ribs number as few as 14 per whorl mit, such specimens (®g. 24) are up to 360 with strong, persistent umbilical bullae. Ribs mm in diameter. These correspond to the ro- are blunt, low, coarse, and prorsiradiate, and bust juveniles in the South Dakota fauna and bear large, conical inner ventrolateral tuber- we tentatively interpret them as macroconchs. cles that are connected by broad prorsiradiate In some there is a long, low siphonal clavus ribs to strong, outer ventrolateral clavi. The corresponding to each ventrolateral horn, suc- siphonal keel is coarse with clavi that cor- cessive clavi being linked by a siphonal ridge. respond in number to that of the outer ven- Low riblets arise from the adapical and ada- trolateral clavi. The largest observed individ- pertural terminations of each ventrolateral ual of the robust type is USNM 252804, 145 horn and loop to the siphonal clavus. In oth- mm in diameter (Cobban and Hook, 1979; ers, small siphonal clavi may be present in the pl. 3, ®gs. 13, 14). Other fragments are still interspaces between larger ones. septate at 65 mm whorl height. European specimens referred to C. wooll- The collections also contain fragments of gari regulare (®gs. 26±32) reach comparable body chambers that show distant ribs with sizes and show the same ontogenetic changes bullae that have migrated out to an inner as the U.S. material described here. ¯ank position, and enlargement of the inner Sutures moderately incised with broad, bi-

ventrolateral tubercles into long, ®nger-like ®d E/L, narrower L, and L/U2 broader than horns (®g. 20B±D), into which the outer ven- U2 (®g. 33). Several sutures were illustrated trolateral clavi are progressively assimilated. by Meek (1876a: pl. 7, ®g. 1e, h) and Mat- The mature body chamber morphology of the sumoto (1965: ®g. 6A, D, F). gracile form is best shown by specimens DISCUSSION: Gracile microconchs and ro- from the Chispa Summit Formation at Chis- bust macroconchs can be recognized from an pa Summit in Jeff Davis County, Trans-Pe- early ontogenetic stage; the former dominate cos Texas. Here, compressed adult body the Black Hills assemblages upon which the chambers (Kennedy et al., 1989: ®g. 23) that above description is largely based. Haas overlap in size and ornament with the simi- (1946) analyzed variation in C. woollgari,in larly compressed juveniles from the Carlile large part on the basis of 325 specimens from Shale of Wyoming already described reach a single concretion he believed to have been diameters of up to 200 mm. The intercostal collected by the Powell Survey from south- whorl section is compressed oval. Distant ern Utah. We have never seen such concre- ribs bear ¯ared umbilical bullae, which pro- tion preservation from Utah and suspect rath- gressively migrate out from the umbilical er that the material is from the Black Hills. shoulder; each ventrolateral horn is linked by Haas's Forma typica (1946: pl. 13, ®gs. 1, a high rib to a siphonal clavus. The costal 4±19; text-®gs. 3±5, 25±29, 31±38) are ma- whorl section is markedly fastigiate and a croconch fragments; the original of his plate low siphonal ridge links the clavi. The ®nal 14, ®gure 11, seems to be a crushed Plesia- portion of the body chamber before the adult canthoceras wyomingense Reagan, 1924. Va- aperture is smooth and tubular without major riety regularis, the name selected for the late ribs, but sometimes bears nontuberculate lir- subspecies of woollgari (Haas, 1946: 154, pl.

← Fig. 40. Collignoniceras praecox (Haas, 1946). A±C. USNM 498273, gracile form from locality 11; D, E. Cast of holotype, USNM 318219 (formerly 103913), robust form; F. Cast of paratype USNM 103913d, gracile form; G. Cast of paratype USNM 103913a; H. Cast of paratype USNM 103913b, gracile form; I. Cast of paratype USNM 103913e, robust form, all from the Carlile Shale, C. praecox Zone, at USGS Mesozoic locality 18872, 3.2 km southeast of Fairburn, Custer County, Wyoming. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_52 Allen Press • DTPro System File # 01235TQ

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Fig. 41. Collignoniceras praecox (Haas, 1946), gracile form. A, B. USNM 498257; C, D. USNM 498256; E, F. USNM 498274; G, H. USNM 498275; I, J. USNM 498276; K±M. USNM 498277; N, O. USNM 498278; P, Q. USNM 498279; R, S. USNM 498280, T, U. USNM 498281; V, W. USNM BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_53 Allen Press • DTPro System File # 01235TQ

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Fig. 42. Collignoniceras praecox (Haas, 1946), gracile form. USNM 498267 from locality 15. Fig- ures are ϫ1.

16, ®gs. 1±17; text-®gs. 10±12, 59±74, 78, 15, ®gs. 1±6, 9, 10; text-®gs. 7±9, 40±44, 80, 81, 83), is based on gracile juvenile mi- 46±58) and variety tenuicostata (Haas, 1946: croconchs. Variety crassa (Haas, 1946: 153, 155, pl. 16, ®gs. 18±21; text-®gs. 13, 14, 75± pl. 14, ®gs. 1, 5; pl. 15, ®gs. 7, 8; text-®gs. 77, 82) are based on juvenile microconchs. 6, 39) is based on evolute, robust, juvenile C. woollgari var. praecox of Haas (1946: microconchs. Variety intermedia (Haas: 155, pl. 16; ®gs. 2±33: pl. 17, ®gs. 1±5; pl. 1946: 154, pl. 14, ®gs. 2±4, 6±10, 12±16; pl. 18, ®gs. 1, 8, 9; text-®gs. 15±18, 79, 84±90)

←

498282, all from locality 15. X, Y. USNM 498283, from locality 12. Z±B1. USNM 498284 from locality 19. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_54 Allen Press • DTPro System File # 01235TQ

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Fig. 44. Collignoniceras praecox (Haas, 1946), robust form. S. Jorgensen Collection, no. Co1087A, from 6±9 m (20±30 ft) below base of Turner Sandy Member, SW¼ sec. 25, T. 7S, R. 6E, Fall River County, South Dakota. Figures are ϫ1.

is based on specimens from USGS localities his other varieties. As already noted, subse- 12642 and 18872 in the Black Hills of South quent collecting shows that this form, here Dakota, and Haas speci®cally noted that the regarded as a fully separate species (C. prae- variety is unknown from any other locality cox), is younger than C. woollgari regulare. in the Black Hills area or in the block from Collignoniceras praecox differs in the ``southern Utah'' that yielded specimens of marked differentiation of ribs into strong and

← Fig. 43. Collignoniceras praecox (Haas, 1946). A, B. USNM 498258; C, D. USNM 498285; E, F. USNM 498263, all robust forms from locality 15. G. S. Jorgensen Collection no. Co388A, robust form from the lower part of the Carlile Shale, sec. 25, T. 7S, R. 6E, Fall River County, South Dakota. H. USNM 498286, gracile forms from locality 14. I, J. USNM 299167, robust form from locality 21. K± M. USNM 498266, robust form from locality 15. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_56 Allen Press • DTPro System File # 01235TQ

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Fig. 45. Collignoniceras praecox (Haas, 1946), robust form. BHI 2101, Carlile Shale, C. praecox Zone, center of sec. 35, T. 8S, R. 1E, Fall River County, South Dakota. Figure is ϫ1.

bullate and weak and nonbullate in the ear- primary ribs, many ventrolateral riblets, and liest stages of development, persistence of an entire or minutely crenulate keel. differentiated ribbing through middle Matsumoto (1965: 11) also analyzed var- growth, and the development in the adult of iation in juvenile Collignoniceras woollgari inner ventrolateral bullae or spines on distant regulare from a collection made from 10 m BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_57 Allen Press • DTPro System File # 01235TQ

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Fig. 47. Collignoniceras praecox (Haas, 1946). External suture of paratype USNM 103913e, copy of Haas, 1946, ®g. 87. Specimen is from the Middle Turonian C. praecox Zone, lower part of Carlile Shale, USGS Mesozoic locality 18872, near Fairburn, South Dakota.

dle and later phragmocone whorls of the latter, there are more siphonal tubercles than ventrolateral ones, and looped ribs connect opposite ventrolateral horns. This generally also separates body chambers, but multiple siphonal tubercles sometimes appear on adult body chambers of C. woollgari regulare,as do looped ribs. Collignoniceras vermilionense (Meek and Hayden, 1860) (see below) is a much smaller species, showing comparable dimorphism to that of C. woollgari regulare. Inner whorls are coarsely ribbed. In the microconch, which is adult at 55 mm diameter, coiling is very evolute. There are only outer ventrolateral clavi whereas in C. woollgari regulare of this size, there are both inner and outer ventrolateral clavi. OCCURRENCE: Widely distributed in the Western Interior of the United States from northern Montana south to Trans-Pecos Texas, and from central Utah east to western Iowa. It is also found in northeastern Texas as well Fig. 46. Collignoniceras praecox (Haas, as California and Oregon. The subspecies also 1946), robust form. BHI 2101, Carlile Shale, C. occurs in Canada (Manitoba), northern Mex- praecox Zone, center of sec. 35, T. 8S, R. 1E, Fall River County, South Dakota. Figure is ϫ1. ico (Chihuahua), southern England (®gs. 27A±G) and France (®gs. 26, 28±32).

above the base of the Carlile Shale 3.2 mi Collignoniceras vermilionense (Meek and southwest of Newcastle, Wyoming. His Hayden, 1860) groups A through D and F are probably all Figures 34D±H, 35A, 37 (pars) juvenile microconchs, his group E, juvenile Ammonites vermilionensis Meek and Hayden, macroconchs. 1860: 177. Collignoniceras woollgari regulare differs Mortoniceras? vermilionense (Meek and Hay- from the nominate subspecies in that in mid- den). Meek, 1876a: 450, pl. 7, ®g. 2. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_58 Allen Press • DTPro System File # 01235TQ

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TABLE 5 Dimensions of Collignoniceras praecox (Haas, 1946)a

Mortoniceras vermilionense (Meek and Hayden). 26 primary ribs per whorl visible at a di- Stanton, 1894: 180, pl. 44, ®g. 1. ameter of approximately 15 mm. All arise at Mortoniceras vermilionense (Meek and Hayden). the umbilical seam and strengthen across the Logan, 1898: 472, pl. 104, ®g. 1. umbilical shoulder. At the smallest observed Prionotropis vermilionensis (Meek and Hayden). Reeside, 1927: 10. diameter, there is a slight differentiation into Collignoniceras vermilionense (Meek and Hay- stronger ribs with umbilical bullae separated den). Cobban, 1983: 17, pl. 5, ®gs. 38±40 (non by somewhat weaker ribs with weak or no 41, 42, ϭ C. praecox). bullae, but this differentiation declines and [Collignoniceras vermilionense]. Ludvigson et al., disappears with increasing size, and the ribs 1994: ®g. 7a. coarsen progressively. All ribs are blunt and recti- to feebly prorsiradiate on the ¯anks. TYPE: Holotype by monotypy is USNM 224 (®g. 34D, E), from a limestone concre- Each rib strengthens into an inner ventrolat- tion in the Carlile Shale, inferred to be the eral bulla from which the rib projects strong- middle Turonian Collignoniceras praecox ly forward, declining markedly in strength Zone, on the Missouri River opposite the and bearing a weak outer ventrolateral cla- mouth of the Vermillion (formerly Vermil- vus. ion) River in Dixon County, Nebraska. USNM 299166 (Cobban, 1983: pl. 5, DIAGNOSIS: A fairly small, keeled, evolute ®gs. 39, 40; ®g. 34F, G), 56.0 mm in di- species that has closely spaced primary and ameter, with inner whorls that are essen- secondary ribs on the innermost whorls, but tially identical to those of the holotype mostly widely spaced, single, prorsiradiate shows the succeeding ontogenetic stages. ribs on later whorls. Ribs arise from low um- Coiling is very evolute, with U/D ϭ 0.46 bilical bullae and terminate in arcuate ven- (table 3); the umbilicus is shallow, with a trolateral swellings that may or may not sup- low wall. The intercostal whorl section is port inner ventrolateral tubercles and faint very compressed, with a whorl breadth to outer ventrolateral clavi. Keel is moderately height ratio of 0.63, the greatest breadth high and strengthened into asymmetric clavi just outside the umbilical shoulder. The corresponding to the ribs. costal ratio is 0.76, the greatest breadth at DESCRIPTION: The holotype is a juvenile. the umbilical bulla. The inner whorls are Coiling is very evolute, with U/D ϭ 0.44± densely and evenly ribbed with 12 primary 0.46 (table 3). The umbilicus is shallow, with ribs per half whorl. There are a total of 18 a low, ¯attened wall and broadly rounded ribs on the outer whorl, all bullate prima- shoulder. The ¯anks are ¯attened. There are ries. They are narrow, very distant, prorsi- BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_59 Allen Press • DTPro System File # 01235TQ

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Fig. 48. Collignoniceras percarinatum (Hall and Meek, 1856). A, B. USNM 498287; C, D. USNM 498288; E, F. USNM 498289; G, H. USNM 498290, all from locality 15. I. Paralectotype AMNH 46057; J. Copy of Hall and Meek's original ®gure of AMNH 31995 (1856: pl. 4, ®g. 2a). K, L. USNM 498291; M, N. USNM 498292; O, P. USNM 498293, all from locality 15. Q. Paratype AMNH 46058; R±U, V. lectotypes. R is a copy of the original ®gure of Hall and Meek (1856: pl. 4, ®g. 2b) (AMNH 31994), U is AMNH 31994, and V is AMNH 30036, part and counterpart of the same specimen. All of the types (I, J, Q, R, U, V) are from rocks now assigned to the middle part of the Carlile Shale, ``Five miles below the mouth of Vermilion River on the Missouri'', that is to say, probably from the northeast side of Volcano Hill in sec. 3, T. 31N, R. 5E, Dixon County, Nebraska. S, T. USNM 299165 from locality 21. Figures A±I, K±P are ϫ2; ®g. J, Q±V are ϫ1.

radiate to the ventrolateral shoulder, where has one side preserved. This specimen, 83.8 each bears a pronounced bullate ventrolat- mm in diameter, has a body chamber occu- eral tubercle at the smallest diameter visi- pying the last one-half whorl and terminat- ble, but which thereafter declines and ef- ing in a pronounced rostrum 35 mm long. faces, leaving a weakening rib that sweeps At the smallest diameter visible, approxi- forward and is markedly concave over the mately 7.5 mm, ribs are single and of equal ventrolateral shoulder. There is a high si- height. All ribs at larger diameters are pri- phonal keel with long siphonal clavi, cor- mary, narrow, and prorsiradiate. They num- responding in number to, but displaced ber 32 per whorl at a diameter of about 11 adaperturally of, the ventrolateral clavi or mm, 26 per whorl at about 19 mm diameter, their site. The umbilical bullae decline and 24 per whorl at 83 mm diameter. All markedly towards the aperture, suggesting arise from low, inconspicuous bullae on the that this specimen may be close to maturity. narrowly rounded umbilical shoulder, in- A magni®cent specimen from a limestone cline forward on crossing the ¯ank, rise into concretion from the Carlile Shale near low bullae on the ventrolateral shoulder, and Yankton, South Dakota, in the S. D. Jorgen- then bend forward sharply and disappear sen collection (Co1970A: ®g. 34H, table 3) close to the keel. Bullae are reduced in size BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_60 Allen Press • DTPro System File # 01235TQ

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Fig. 49. A±K, M±T. Collignonicerites collisniger n. sp. A, B. paratype USNM 498295; C, D. paratype USNM 498296; E, F. paratype USNM 498297; G±I. paratype USNM 498298, J. paratype BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_61 Allen Press • DTPro System File # 01235TQ

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TABLE 6 Dimensions of Collignonicerites collisniger n. gen., n. sp.a

near the aperture, and the last two septa are 498250 (®g. 34A±C) is from the Carlile approximated. Shale at USGS Mesozoic locality D12683 The suture, illustrated by Meek (1876a: pl. (locality 13 in ®g. 1) in the NE1/4 NW1/4 7, ®g. 2b), is fairly simple and proportioned sec. 32, T. 94 N, R. 50 W, Union County, like that of the juvenile Collignoniceras South Dakota. woollgari ®gured by Meek (1876a: pl. 7, ®g. DIAGNOSIS: A rather small, evolute species 1e). Part of a suture near the aperture of the that has prorsiradiate primary and secondary large Jorgensen specimen is shown in ®g. ribs throughout. Primary ribs bear umbilical 35A. bullae and inner and outer ventrolateral tu- DISCUSSION: This is a very rare species. bercles. Keel supports a clavate tubercle for Cobban (1983) initially considered it Col- each rib. Primary ribs are commonly ¯ared. lignoniceras praecox (Haas, 1946). The lat- DESCRIPTION: The holotype (®g. 36A) is a ter, however, has straight, rectiradiate ribs small adult, 38.3 mm in diameter, embedded that terminate in sharp horns in contrast to in part of a limestone concretion, so that only the prorsiradiate ribs of C. vermilionense one side is visible. Slightly more than one that bear fowardly inclined ventrolateral half of the last whorl is body chamber, fol- bullae. lowed by a long rostrum. The umbilicus is OCCURRENCE: Known only from the Car- shallow with U/D ϭ 0.42 (table 4). The um- lile Shale in northeastern Nebraska and bilical shoulder is narrowly rounded. Whorls southeastern and southwestern South Dakota; are apparently higher than wide, are some- Collignoniceras praecox Zone. what ¯attened, and have a narrowly rounded ventrolateral shoulder. The keel is fairly high Collignoniceras jorgenseni, new species and accentuated into clavi that correspond to Figures 34A±C, 35B, 36, 37 (pars), 38 (pars) the ribs. Those clavi corresponding to the DERIVATION OF NAME: For Steven D. Jor- secondary ribs are low and symmetric; those gensen of Omaha, Nebraska, who donated corresponding to the primary ribs are high the holotype and several other specimens and asymmetric with the steep slope forward. from his extensive collections of Cretaceous Ornament of the holotype consists of pror- ammonites. siradiate ribs of unequal heights and umbil- TYPES: Holotype is USNM 498249 (®g. ical and ventrolateral tubercles as well as si- 36A) from the Collignoniceras praecox Zone phonal clavi. All ribs are narrow, and all in the middle Turonian Carlile Shale at arise on the umbilical shoulder. Primary ribs USGS Mesozoic locality D13834 in the arise from narrow, umbilical bullae or from SW1/4 SW1/4 sec. 25, T. 7 S, R. 6 E, Fall narrow, bullate tubercles. Each primary is in- River County, South Dakota. Paratypes clined forward on crossing the ¯ank and then USNM 498253 (®gs. 35B, 36B±D, K) rises into a prominent, arcuate ventrolateral 498254 (®g. 36H±J), 498251, and 498252 swelling that supports a weak inner ventro- are from the same locality. Paratype USNM lateral tubercle and a weaker outer ventrolat-

← USNM 498299; K. paratype USNM 498300; M, R±T. holotype, USNM 498294; N. O±Q. paratype USNM 498302. L. Collignoniceras woollgari regulare (Haas, 1946), robust form. All specimens are from locality 12. Figures A±L, O±T are ϫ2; ®gures M, N are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_62 Allen Press • DTPro System File # 01235TQ

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is separated by one to four low, narrow, secondary ribs that are usually accentuated on the ventrolateral shoulder, but do not bear tubercles. Total primary and secondary rib counts per whorl are as follows: 35 at a diameter of approximately 6 mm, 38 at a diameter of about 14 mm, and 38 at a diameter of about 22.5 mm. That the holotype is a small adult is re- vealed by the weakening and disappearance of tubercles on the last part of the body chamber, and by the presence of a long (13.5 mm) rostrum. Only short sections of the suture are visible. Two paratypes (®g. 36B±D, K and 36H± J) from the same concretion as that of the holotype (®g. 36A) are adults 61 and 65 mm in diameter with umbilical to shell diameter ratios of 0.38 and 0.45, respectively. Both have crushed body chambers with the rostra broken off; the phragmocones are slender and probably somewhat crushed. Ribbing is sparser than that of the holotype, and primaries and secondaries alternate. Another adult paratype (un®gured) from the same concretion has a diameter of 50.5 mm and an umbilical to shell diameter ratio of 0.48. Its body chamber is also crushed, but the prominent rostrum is preserved. Most primary ribs are separated by two secondaries. The holotype (®g. 36A) and paratypes USNM 498250 (®g. 34A±C) and 498252 Fig. 50. Prionocyclus hyatti (Stanton, 1894). represent the robust form of the species. Par- A, B. Paralectotype USNM 22941a. C±E. lecto- atypes USNM 498253 (®g. 36B±D, K) and type, USNM 22941. F±H. Paralectotype USNM 498254 (®gs. 36H±J) represent the gracile 22941b. All specimens are from the Middle Tu- form. ronian P. hyatti Zone, Codell Sandstone (Pugnel- A complete suture is not visible on any lus Sandstone) of Williams Creek and Poison specimen. The lateral lobe and adjoining sad- Canyon, Huerfano Park, Colorado. All ®gures are dles on one of the paratypes is shown in ®g. ϫ1. 35B. DISCUSSION: Collignoniceras jorgenseni eral tubercle before diminishing and disap- resembles C. vermilionense in size, in its pearing at an acute angle to the keel. Some wide umbilicus, and in its conspicuous pror- primaries are ¯ared. There are 12 primaries siradiate ribbing, but C. jorgenseni differs in on the outer whorl. Each pair of primary ribs its dense ribbing and abundant secondaries.

→ Fig. 51. Prionocyclus hyatti (Stanton, 1894). A±C. USNM 498315; D, E. USNM 498316; F, G. USNM 498303; H, I. USNM 498317; J±L. USNM 498318, all gracile forms from locality 41. M±O. USNM 498310, gracile form from locality 40. P, Q. USNM 498319, gracile form from locality 23. R. USNM 498320, gracile form from locality 41. S±U. USNM 498306, V, W. USNM 498321, both gracile forms, both from locality 41. X, Y. USNM 498311, gracile form from locality 40. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_63 Allen Press • DTPro System File # 01235TQ

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Both forms occur with C. praecox and C. ribs and weak intercalaries that give these percarinatum. small specimens a highly distinctive appear- OCCURRENCE: Known only from the lower ance. There are up to 10 ¯ared primary ribs part of the Carlile Shale in the Black Hills per whorl with well-developed umbilical bul- area of western South Dakota, Collignoni- lae. The primary ribs are narrow, high, and ceras praecox Zone. prorsiradiate on the ¯anks, and each bears a well-developed feebly clavate inner ventro- Collignoniceras praecox (Haas, 1946) lateral tubercle from which the rib projects Figures 39±47 sharply forward to a clavate outer ventrolat- Prionotropis woollgari Meek (non Mantell) var. eral tubercle. Between the ¯ared primary ribs praecox Haas, 1946: 155, pl. 16, ®gs. 22, 23; are up to four much weaker, nonbullate ribs pl. 17, ®gs. 1±5; pl. 18, ®gs. 1, 8, 9; text-®gs. without ventrolateral tubercles. These speci- 15±18, 79, 84±90. mens closely match one of the paratypes Collignoniceras vermilionense (Meek and Hay- (USNM 103913b, Haas, 1946: pl. 16, ®g. 29, den). Cobban, 1983: 17 (pars), pl. 5, ®gs. 41, cast re®gured here as ®g. 40H), which shows 42. similar ornament to a whorl height of 14 mm, Subprionocyclus percarinatus (Hall and Meek). Cobban, 1983: 18 (pars), pl. 5, ®gs. 26±29, 32, at which point each weaker rib bears a feeble 33, 36, 37. inner ventrolateral tubercle with pairs of both Collignoniceras praecox (Haas, 1946). Reyment inner and outer ventrolateral tubercles pre- and Kennedy, 2001: ®g. 1 k±n. sent on the stronger ribs. Beyond this whorl height, the weaker ribs efface markedly, TYPES: Holotype is USNM 318219 (®g. leaving primary ribs and lirae on the ¯anks 40D, E) (formerly 103913), paratypes (USNM 498283, ®g. 41X, Y). In other spec- USNM 103913a±e (®g. 40F±I; USNM 103913C not ®gured), all from the Carlile imens (®g. 39G±K) this stage of markedly Shale, middle Turonian Collignoniceras differentiated ribs is followed, to a diameter praecox Zone at USGS Mesozoic locality of at least 35 mm, by a stage with crowded, 18872, 3.2 km southeast of Fairburn, Custer dense bullate and nonbullate primary and oc- County, South Dakota. casional intercalated ribs that total up to 55 DIAGNOSIS: Moderate to fairly large spe- per whorl. These specimens are very similar cies with highly variable juvenile whorls that to paratype USNM 103913d (Haas, 1946: pl. range from densely ribbed with distinct pri- 18, ®gs. 8, 9; ®g. 40F); the ribs are straight maries and secondaries to more evenly and and prorsiradiate, and are separated by nar- sparsly ribbed. Adults have sparser and usu- row interspaces. Each rib bears a bullate in- ally weaker ribs and prominent ventrolateral ner ventrolateral tubercle from which the rib horns. projects abruptly forward and links to a cla- DESCRIPTION: The present collections are vate outer ventrolateral tubercle. All these nearly all juveniles of the gracile form. Coil- variants have a pronounced siphonal keel ing is fairly evolute, with an umbilicus to with numerous siphonal clavi that are either shell diameter ratio of just under 0.30 (table as numerous or fewer in number per whorl 5). The whorl section is compressed oval in than the ventrolateral tubercles. In middle intercostal section and polygonal in costal growth there is an abrupt change to widely section with whorl breadth to height ratios of spaced ribs (®gs. 39X±Z; 40D, E, I; 41X, Y). as little as 0.62. Ribbing is highly variable at Bullate primaries alternate with nonbullate small diameters (®gs. 39, 40A±C, R±H; long and short ribs, all with conical inner and 41A±W), but generally shows a marked dif- clavate outer ventrolateral tubercles, the ferentiation into strong and weak primary compressed whorl section and feeble ¯ank

← Fig. 52. Prionocyclus hyatti (Stanton, 1894). A±C. USNM 498309; D±F. USNM 498322; G, H. USNM 356904; I, J. USNM 498323; K, L. USNM 498324, all gracile forms from locality 41. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_66 Allen Press • DTPro System File # 01235TQ

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Fig. 53. Prionocyclus hyatti (Stanton, 1894). USNM 498312, gracile form from locality 41. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_67 Allen Press • DTPro System File # 01235TQ

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Fig. 54. Prionocyclus hyatti (Stanton, 1894). USNM 498325, gracile form, from locality 41. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_68 Allen Press • DTPro System File # 01235TQ

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Fig. 55. Prionocyclus hyatti (Stanton, 1894). A, B. USNM 498326, gracile form from locality 41; C, D. USNM 498304, robust form from locality 40. E±G. USNM 498327, gracile form from locality 42. H±J. USNM 498328, gracile form from locality 41. All ®gures are ϫ1.

ornament being distinctive at this size; the intersect the line of the keel at an acute angle, holotype is just such a specimen (®g. 40D, with the interspaces between ribs covered in E). In ventral view these specimens show the growth striae, lirae, and riblets. At this stage, ribs extending in attenuated form forward to the crenulations on the keel are very sub-

→ Fig. 56. Prionocyclus hyatti (Stanton, 1894). A, B. USNM 498329, robust form from locality 41. C, D. USNM 498305, robust form from locality 42. E±G. USNM 498308, robust form from locality 40. H±J. USNM 498307, robust form from locality 41. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_69 Allen Press • DTPro System File # 01235TQ

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Fig. 57. Prionocyclus hyatti (Stanton, 1894). A, B. USNM 498330; C, D. USNM 498331, both robust forms from locality 23. All ®gures are ϫ1.

dued. In some there are merely irregular el- have sparse primary ribs ending in pro- evations; in others (e.g., Haas, 1946: pl. 17, nounced ventrolateral tubercles or horns. ®g. 4; ®g. 40I) the keel is almost entire. Our Several sutures were shown by Haas largest specimen of this kind, USNM 498267 (1946: ®gs. 79, 84±90); that of one of the (®g. 42), shows progressive effacement of paratypes (USNM 103913e) is reproduced ¯ank ornament and the coalescence of inner here (®g. 47). and outer ventrolateral tubercles into blunt DISCUSSION: Haas (1946: 156) rightly re- inner ventrolateral horns. USNM 498284 marked on the distinctive features of this (®g. 41Z±BЈ) shows the same transition at species: ``the accelerated development of only 60 mm diameter. this variety from maturity to the latest on- Juveniles of the robust form are evolute togenetic stage has been alluded to in its with U/D up to 0.35 (®g. 43E, F; table 5). name. The differentiation between stronger Their innermost whorls may or may not and weaker ribs and the later disappearance show differentiation into strong, bullate and of those of the latter kind cause this variety, weaker, nonbullate ribs. In middle growth, of all of the various forms of P. woollgari, the few available specimens have strong, dis- most to resemble the genus Prionocyclus. tant primary ribs with bullae and inner and Furthermore it is the only form of, or close outer ventrolateral tubercles differentiated to to P. woollgari here dealt with that might whorl heights of up to 30 mm, with shorter, perhaps be granted the status of an indepen- weaker ribs between. Adults (®gs. 44±46) dant species.'' Collignoniceras praecox thus BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_71 Allen Press • DTPro System File # 01235TQ

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TABLE 7 Dimensions of Prionocyclus hyatti (Stanton, 1894)a

Fig. 58. Prionocyclus hyatti (Stanton, 1894). USNM 487697, robust form from locality 23. Figures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_72 Allen Press • DTPro System File # 01235TQ

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nile C. woollgari woollgari and C. praecox, but the strong siphonal clavi of the former, corresponding to the outer ventrolateral tubercles in number, separates them; adults are very different. OCCURRENCE: Known with certainty only from the lower part of the middle Turonian Carlile Shale of South Dakota, Collignoni- ceras praecox Zone.

Collignoniceras percarinatum (Hall and Meek, 1856) Figure 48 Ammonites percarinatus Hall and Meek, 1856: 396, pl. 4, ®g. 2a, b. Prionotropis woolgari (sic) Mantell (sp.). Stanton, 1894: 174. Ammonites percarinatus Hall and Meek. Diener, 1925: 32. Prionotropis percarinatus (Hall and Meek). Haas, 1946: 150, 158. Collignoniceras woollgari (Mantell). Stephenson, 1953: pl. 12, ®g. 3. Subprionocyclus percarinatus (Hall and Meek). Cobban, 1983: 18, pl. 5, ®gs. 30, 31, 34, 35 (non ®gs. 26±29, 32, 33, 36, 37 (ϭ Collignon- Fig. 59. Prionocyclus hyatti (Stanton, 1894). iceras praecox)); pl. 15, ®gs. 5, 6. USNM 498332, detached ventrolateral horn, from Prionocyclus percarinatus (Hall and Meek). Cob- locality 23. Figures are ϫ1. ban, 1984a: 75, 84. non Prionocyclus percarinatus (Hall and Meek, differs from C. woollgari regulare in the 1856). Kirkland, 1996: 99, pl. 40, ®gs. B±F, I. middle growth stages by the persistence of TYPES: Lectotype AMNH 31995 (®g. 48J), long and short ribs, with ventrolateral tu- designated by Cobban (1983: 18), and para- bercles outnumbering umbilicals. The pres- types are from ``Five miles below the mouth ence of a near continuous keel where C. of Vermilion River on the Missouri'' (Hall woollgari regulare has clavi equal in num- and Meek, 1856: 396). As noted by Cobban ber to the ventrolateral tubercles is also dis- (1983: 18), the specimens came from rocks tinctive. Adult C. woollgari are much larger now assigned to the middle Turonian Carlile than those of C. praecox, and have a highly Shale, probably from the northeast side of distinctive ribbing and ventral ornament. Volcano Hill in sec. 3, T. 31 N, R. 5 E, Dixon There are closer similarities between juve- County, Nebraska.

Fig. 60. Suture of Prionocyclus hyatti (Stanton, 1894). Copy of Haas, 1946, ®gure 45. The original is from the Blue Hill Member of the Carlile Shale of Kansas. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_73 Allen Press • DTPro System File # 01235TQ

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Fig. 61. Distribution of Prionocyclus hyatti (Stanton, 1894) in the Western Interior Seaway during the early Middle Turonian. Land areas indicated by stipple. Modi®ed after Cobban et al. (1994). BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_74 Allen Press • DTPro System File # 01235TQ

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Fig. 62. Prionocyclus albinus (Fritsch, 1872). A. USNM 498333, from locality 32. B. USNM 220385, from locality 22. C±E. USNM 498334, from USGS Mesozoic locality 35. F. USNM 498335, from locality 63. G, H. USNM 498336, from locality 32. I±K. USNM 356899, from locality 40. L, M. USNM 498337, from locality 63. N, O. USNM 498338, from locality 32. P. USNM 498339, from locality 32. All ®gures are ϫ1.

DIAGNOSIS: A small species that has close- DESCRIPTION: Hall and Meek's descrip- ly spaced, prorsiradiate ribs of rather uniform tion is as follows: ``Discoidal, depressed; height. Ribs bend forward at the ventrolateral umbilicus wide and shallow; volutions shoulder and fade out at the low keel. Keel about four or ®ve, all visible in the umbi- notched by low clavi that correspond in num- licus, scarcely one fourth of each embraced ber to the ribs. in the succeeding one; shell thin; surface BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_75 Allen Press • DTPro System File # 01235TQ

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Fig. 63. Prionocyclus macombi Meek, 1876. A, B. lectotype, USNM 20259, from the Juana Lopez Member of the Carlile Shale in Colfax County, New Mexico. C±G. USNM 22940, the originals of Stanton, 1894, plate 41, ®gures 1±3, from ``near Mancos, southwestern Colorado, about 400 feet above the base of the Colorado Cretaceous shales, where it is associated with Ostrea lugubris and Inoceramus dimidius.'' Figures A±C, E±G are ϫ1; ®gure D is ϫ2.

marked by thirty-eight to forty-®ve promi- ty, South Dakota, seem assignable to this nent ¯exuous sharp ribs some of which species (®g. 48A±H, K±P). The specimens originate in the umbilicus, and others upon are fairly evolute with umbilical to shell di- the lateroventral margin, and all extend to ameter ratios of 0.33±0.48 and 32±38 ribs on the dorsolateral edge, where they bend the outer whorl. Ribs are prorsiradiate, arise abruptly forward, and terminate before at the umbilicus, cross the ¯ank, and curve reaching the dorsal line, which is marked forward at the ventrolateral shoulder, where by a thin sharp carina extending to the ap- they are accentuated a little or actually form erture. Ribs thickened and sometimes no- low clavate tubercles. The low keel bears dose towards the periphery. Our specimens asymmetrical clavi that have the steeper are all casts of the interior with fragments slope on the adoral side. Each siphonal cla- of the shell adhering, and the condition is vus matches a rib. such as to give no means of determining DISCUSSION: The presence of a siphonal the character of the septa.'' clavus for each rib supports the assignment A suite of small specimens from limestone to Collignoniceras rather than to Prionocy- concretions in the lower part of the Carlile clus. In later whorls, the species resembles Shale at locality 15 in ®gure 1, USGS Me- C. vermilionense in having single prorsira- sozoic locality D10697, in Fall River Coun- diate ribs that bend forwards sharply on the BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_76 Allen Press • DTPro System File # 01235TQ

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Fig. 64. Prionocyclus macombi Meek, 1876. A, B. USNM 498353; C. USNM 498354; D±F. USNM 498355; G. USNM 498356; H. USNM 498357; I. USNM 498358; J±L. USNM 498359; M±O. USNM 498360, all variants of the gracile form, from locality 27. All ®gures are ϫ1.

ventrolateral shoulder. The latter species, TYPE SPECIES: Collignonicerites collisniger however, is much more coarsely ribbed and n. gen., n. sp. has larger, more pronounced siphonal clavi. DIAGNOSIS: Small, adult at 18 mm or less, OCCURRENCE: Known only from the mid- involute with compressed whorl section. Dis- dle Turonian Carlile Shale, Collignoniceras tant, blunt umbilical bullae give rise to pairs praecox Zone, of southeastern South Dakota, or groups of ribs, with one or several non- northeastern Nebraska, and western Minne- bullate long ribs between; ribs blunt, crowded, sota. Specimens ®gured as Prionocyclus per- prorsiradiate, ¯exuous, all with small outer carinatus by Kirkland (1996: 99, pl. 40, ®gs. ventrolateral clavi. Venter broad, ¯at, with B±F, I) from the Collignoniceras woollgari strong siphonal keel and weak siphonal clavi regulare Subzone in the Mancos Shale of equal in number to ventrolateral clavi but dis- northeastern Arizona have differentiated ribs placed adaperturally on phragmocone. Orna- to a much larger size than on C. percarinatus ment declines on adult body chamber, which and represent some other species. Likewise, may be constricted; umbilical bullae present, specimens recorded as Subprionocyclus per- ¯ank ribs decline but ventrolateral ornament carinatus? from the Mancos Shale of west- persists; siphonal clavi become irregular. central New Mexico by Hook et al. (1983: The suture is very simple with broad bi®d sheet 1) also belong to some other species. E/L with few incisions, narrower L, and

nearly entire L/U2. Genus Collignonicerites, new genus DISCUSSION: Decline in ornament and the DERIVATION OF NAME: Collignoniceras development of constrictions indicates that plus ites (Latin)Ðdescendant of Collignoni- the species attains maturity at 17 mm or less, ceras. while the very simple adult suture indicates

→ Fig. 65. Prionocyclus macombi Meek, 1876. A, B. USNM 498361; C, D. USNM 498362, both gracile forms. E±G. USNM 498363, robust form. H±J. USNM 498364; K, L. USNM 498365; M, N. USNM 498366; O±Q. USNM 498367, all gracile forms. R±T. USNM 498368, robust form; U±W. USNM 498369; X, Y. USNM 498370, both gracile forms; Z, A1. USNM 498371, robust form. All specimens are from locality 6. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_77 Allen Press • DTPro System File # 01235TQ

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Fig. 66. Prionocyclus macombi Meek, 1876. USNM 498372, a robust form from locality 6. All ®gures are ϫ1.

that these specimens are more than mere ju- earliest developmental stages of Collignoni- veniles of some larger form. Collignoniceri- ceras with which Collignonicerites collisni- tes resembles Collignoniceras; the very sim- ger gen. et sp. nov. co-occurs. ple suture of Collignoniceras (e.g., Matsu- Collignonicerites is an analog of Priono- moto, 1965: text-®g. 6A) is retained to ma- cyclites Kennedy, 1988, a diminitive genus turity in Collignonicerites. The phragmocone closely related to Prionocyclus. Species of ornament of Collignonicerites with outer the two genera share certain features com- ventrolateral clavi equal in number to si- mon to progenetic dwarfs, including a great- phonal clavi and ribs springing in groups ly simpli®ed suture. However, they differ in from bullae resembles that of the ornamented that species of Prionocyclites are more evo-

→ Fig. 67. Prionocyclus macombi Meek, 1876. A. USNM 420139; B. USNM 498373; C±E. USNM 498374, all gracile forms; F±H. USNM 498375; I±K. USNM 498376; L, M. USNM 498377; N±P. USNM 498378, all robust forms. Q, R. USNM 498379; S, T. USNM 420138; U, V. USNM 420136, all gracile forms. W, X. USNM 498380, robust form. Y, Z. USNM 498381; A1,B1. USNM 498382, both gracile forms. C1,D1. USNM 498383; E1±G1. USNM 498384, both robust forms. A±X, C1±G1 are from locality 36; Y-B1 are from locality 38. Figures A±E are ϫ2; ®gures F±G1 are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_79 Allen Press • DTPro System File # 01235TQ

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lute and lack massive bullae giving rise to The whorl section is compressed, with great- groups of ribs, having instead narrow, dis- est breadth below mid¯ank in intercostal sec- tant, single bullate primaries separated by tion, and shows broadly rounded inner nonbullate primaries and secondaries, an un- ¯anks, ¯attened convergent outer ¯anks, and dulose rather than serrated keel, and a body a relatively broad, ¯attened venter with a chamber with distant feeble bullae, narrow, sharp siphonal keel. The greatest breadth is single ¯ank ribs, and an apertural constric- at the umbilical bullae in costal section. In tion ¯anked by collar-ribs. strongly ornamented variants, blunt umbilical Progenetic species have increasingly been bullae, four to six per whorl, are present in described in recent publications on Late Cre- middle growth. The bullae give rise to two taceous ammonites, as noted by Wright or three primary ribs with up to three non- (1996: xviii; see, for instance, Wright and bullate long ribs separating the bullate Kennedy, 1980; Kennedy, 1988; Kennedy groups. These nonbullate ribs arise either sin- and Cobban, 1990a, 1990b). Dimorphism has gly or in pairs at the umbilical shoulder to been recognized in these dwarfs, demonstrat- give a total of 16 ribs per half whorl. Ribs ing that they are not the microconchs of co- are broad, crowded, ¯exuous, and prorsira- occurring larger taxa. Because of their small diate, and each bears a small outer ventro- size, these dwarf taxa have, in general, only lateral clavus from which the rib projects for- been detected in exceptionally preserved fau- ward and declines in strength, intersecting nas, such as those found in the concretions the line of the siphonal keel in an acute chev- in shale and mudstone facies of the U.S. ron. The keel has even siphonal clavi, equal Western Interior. in number to but displaced adaperturally of OCCURRENCE: Middle Turonian Collignon- the ventrolateral clavi. In feebly ornamented iceras woollgari regulare Subzone of the variants, the umbilical bullae are weak; the Black Hills only. ¯anks bear only striae and lirae on the inner part but the ventrolateral part is similar to Collignonicerites collisniger, new species that already described. The umbilical bullae Figure 49A±K, M±T may decline on the adult body chamber, which may be constricted, while the crenu- DERIVATION OF NAME: collis ϩ niger (Lat- lations on the siphonal keel become irregular. in) black hills, from the type area. The suture is the same as for the genus. TYPES: Holotype is USNM 498294 (®g. DISCUSSION: See under genus. 49M, R±T), paratypes USNM 498295±498302 OCCURRENCE: As for types. (®g. 49A±K, N±Q; 498301 not illustrated), from the middle Turonian Carlile Shale, Col- Genus Prionocyclus Meek, 1876 lignoniceras woollgari regulare Subzone at lo- (ϭ Germariceras Breistroffer, 1947) cality 12 in ®g. 1: USGS Mesozoic locality D13832, Carlile Shale, middle Turonian, C. TYPE SPECIES: Prionocyclus serratocari- woollgari regulare Subzone, sec. 35, T. 8 S, natus Meek, 1871: 298, non Stoliczka, 1865: R. 1 E, Fall River County, South Dakota. pl. 32, ®g. 31, ϭ Prionocyclus wyomingensis DIAGNOSIS: With the characters of the genus. Meek, 1876a: 452. DESCRIPTION: Coiling fairly involute (U/D DIAGNOSIS: Medium to large, adult up to ϭ 0.26) and umbilicus moderately deep (ta- 300 mm in diameter. Size-related dimor- ble 6). The umbilical wall is ¯attened, sub- phism not recognized (due to paucity of vertical, with an evenly rounded shoulder. adults), but the middle growth stages of all

← Fig. 68. Prionocyclus macombi Meek, 1876. A±C. USNM 498385, gracile form from locality 56. D, E. USNM 498386, gracile form from locality 55. F, G. USNM 498387, robust form from locality 56. H, I. USNM 498388, robust form from locality 55. J±L. USNM 498389, gracile form from locality 55. M, N. USNM 498390; O, P. USNM 498391, both robust forms from locality 56. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_82 Allen Press • DTPro System File # 01235TQ

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species occur as involute, relatively com- France (Middlemiss and Moullade, 1968) are pressed, feebly ornamented gracile forms or unsubstantiated. more evolute, less compressed, strongly ornamented, robust forms. Ornament highly Prionocyclus hyatti (Stanton, 1894) variable, typically of stronger primary ribs Figures 50±60 with umbilical, inner, and outer ventrolateral tubercles at some stage in ontogeny separat- Prionotropis hyatti Stanton, 1894: 176, pl. 42, ed by weaker nonbullate and intercalated ribs ®gs. 5±8. that have inner and outer ventrolateral tuber- Prionocyclus hyatti Stanton. Logan, 1898: 468, pl. cles at some stage in ontogeny. Outer ven- 102, ®gs. 5±8. Prionocyclus hyatti Stanton. Grabau and Shimer, trolateral tubercles may decline or not, inner 1910: 228, ®g. 1509e±g. ventrolateral tubercles may bear long septate Prionocyclus hyatti Stanton. Diener, 1925: 156. spines; faint inner lateral tubercles may de- Prionotropis aff. woollgari (Mantell). Moreman, velop during later growth stages. Siphonal 1927: 97 (pars), pl. 13, ®g. 2. keel is strong to weak, and ®nely serrated, Prionotropis aff. P. hyatti Stanton. Adkins, 1928: with the number of serrations exceeding the 250. number of ribs. Prionotropis eaglensis Adkins, 1928: 250, pl. 32, DISCUSSION: The type species of Priono- ®gs. 1, 2. cyclus and Collignoniceras are distinct, es- Pseudaspidoceras eaglense (Adkins). Adkins, pecially in the adult stage, and juveniles can 1931: 53. Prionotropis graysonensis (Shumard). Moreman, be separated because the keel of Collignon- 1942: 213. iceras has siphonal clavi equal in number to Prionocyclus aff. woollgari Meek (not Mantell). that of the ventrolateral tubercles, whereas in Moreman, 1942: 214. Prionocyclus the serrations greatly outnum- Prionotropis woollgari Meek (?non Mantell). ber the ribs (compare ®gs. 17, 18 and ®gs. Haas, 1946: ®g. 45. 78, 84). In Prionocyclus, the marked differ- Collignoniceras hyatti (Stanton). Cobban, 1951: entiation of ribbing present in the early on- 2188, 2191, 2192, ®g. 2. togenetic stages of certain variants of C. Collignoniceras hyatti (Stanton). Cobban and praecox (e.g., ®g. 41F±U) persists to matu- Reeside, 1952: 1018. rity. Collignoniceras hyatti (Stanton). Cobban, 1955: 204, pl. 2, ®g. 10. Germariceras Breistroffer, 1947 (un- Collignoniceras hyatti (Stanton). Hattin, 1962: 79, paged), with Ammonites germari Reuss, pl. 24, ®gs. A±E. 1845 (®g. 109K), from the upper Turonian of Prionocyclus hyatti (Stanton). Powell, 1963: Czech Republic as type species, is best 1220, pl. 166, ®gs. 1, 8±12; text-®g. 5a, c, d, known from the lithographs of Fritsch (1872: f±h. pl. 14, ®gs. 1, 2; pl. 16, ®g. 7), and was re- Collignoniceras hyatti (Stanton). Hattin, 1965: cently revised by Kaplan (1988). It is here ®g. 4 (4). shown to be common and widespread in the Prionocyclus hyatti (Stanton). Matsumoto, 1965: U.S. Western Interior, and shows no charac- 19, pl. 17, ®g. 3. ters that merit separation from Prionocyclus. Prionocyclus hyatti (Stanton). Cobban, 1976: 122, pl. 1, ®g. 1. OCCURRENCE: Middle Turonian of the U.S. Prionocyclus hyatti (Stanton). Kennedy and Cob- Western Interior, Gulf Coast, and northern ban, 1976: pl. 8, ®g. 3. Mexico; upper Turonian of the U.S. Western Prionocyclus hyatti (Stanton). Kauffman, 1977: Interior, Gulf Coast, and northern Mexico, pl. 22, ®g. 1; pl. 26, ®g. 7. Tunisia, Japan, Czech Republic, Germany, Prionocyclus hyatti (Stanton). Hattin, 1977: ®gs. and Kazakhstan. Records from Southeast 8 (9, 11).

← Fig. 69. Prionocyclus macombi Meek, 1876. A±C. USNM 498346, gracile form from locality 39. D, E. USNM 498392, gracile form from locality 36. F, G. USNM 498347, gracile form from locality 39. H, I. USNM 498393, robust form from locality 36. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_84 Allen Press • DTPro System File # 01235TQ

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Prionocyclus hyatti (Stanton). Hattin and Siemers, on outer whorl support umbilical bullae or 1978: ®g. 10. nodate umbilical tubercles. Inner ventrolat- Prionocyclus hyatti (Stanton). Kauffman et al., eral tubercles on adult body chamber may 1978: pl. 5, ®g. 1. enlarge into conspicuous spines. Prionocyclus hyatti (Stanton). Young and Powell, 1978: pl. 1, ®gs. 1±3, 7, 8. DESCRIPTION: The lectotype is a small, ro- Prionocyclus hyatti (Stanton). Merewether et al., bust specimen (Stanton, 1894: pl. 42, ®gs. 5, 1979: pl. 3, ®gs. 1, 2. 6; Matsumoto, 1965: 19, pl. 17, ®g. 3a±c; ®g. Prionocyclus hyatti (Stanton). Cobban, 1984a: 85. 50C±E). Measurements in millimeters and Prionocyclus hyatti (Stanton). Cobban, 1986: ®g. ratios from a plaster cast are as follows: D 3Q. ϭ 31.0, Wb ϭ 14.2 (46%), Wh ϭ 12.0 Prionocyclus hyatti (Stanton). Kennedy, 1988: 75, (39%), Wb:Wh ϭ 1.18, U ϭ 10.6 (34%). Pri- pl. 15, ®gs. 1±24; pl. 16, ®gs. 1±20; pl. 17, ®gs. mary and secondary ribs arise on the umbil- 1±7; text ®gs. 24d, e, f, 25±27, 31b. ical wall and are straight and prorsiradiate on Prionocyclus hyatti (Stanton, 1894). Kennedy and Cobban, 1988: 606, ®gs. 7. 1±8, 11. crossing the ¯anks. They number 29 per Prionocyclus hyatti (Stanton, 1894). Kennedy et whorl at a diameter of approximately 7.5 al., 1989: 89, ®gs. 24f, g, m, 25, 26a, b, 27a±j, mm, and 29 per whorl at a diameter of about s±w. 19 mm. Primary ribs support umbilical bul- Prionocyclus hyatti (Stanton). Cobban and Hook, lae, prominent clavate inner ventrolateral tu- 1989: ®g. 9i, j. bercles, and weak outer ventrolateral clavi. Prionocyclus hyatti (Stanton). Cobban, 1990: pl. Some secondaries bear weaker ventrolateral 5, ®gs. 3±5. tubercles. All ribs bend forward on the ven- Prionocyclus hyatti (Stanton, 1894). Emerson et al., 1994: 210, unnumbered ®gures on pp. 210, trolateral shoulder and fade out at an angle 211. toward the keel. Most primaries and second- Prionocyclus hyatti Stanton, 1893. Kirkland, aries alternate. Toward the adapertural end, 1996: 99, pl. 50, ®gs. L, M. the inner ventrolateral tubercles on the pri- Prionocyclus hyatti (Stanton, 1894). Reyment and mary ribs enlarge and become hornlike, pro- Kennedy, 2001; ®g. 2a±d. jecting outward at a low angle to the venter. TYPES: Lectotype, by subsequent desig- In this manner, the specimen is much like the nation of Matsumoto 1965: 19, is USNM last part of the phragmocone of the remark- 22941, the original of Stanton's 1894: plate able spined individual ®gured by Kennedy 42, ®gures 5, 6 (®g. 50C±E); ®gured para- and Cobban (1976, pl. 8, ®g. 1; ®g. 58) from lectotype is USNM 22941a, the original of the Carlile Shale of Kansas. On the older part Stanton's plate 42, ®gures 7, 8 (®g. 50A, B). of the outer whorl of Stanton's specimen, the They are from the Codell Sandstone (Pug- keel is differentiated into low clavi that cor- nellus Sandstone) of Williams Creek and respond to the ribs, but on the younger part, Poison Canyon, Huerfano Park, Colorado. the clavi seem to number about two per rib. Stanton refers to many specimens from The other specimen ®gured by Stanton Huerfano Park and a few from Coalville, (1894: pl. 42, ®g. 7; ®g. 50A, B) is a small, Utah: all are paralectotypes (e.g., ®g. 50F± gracile, evolute specimen 21.0 mm in diam- H). eter with an umbilical ratio of 49% (mea- DIAGNOSIS: A moderately evolute fairly surements from a plaster cast). The keel is large species ornamented by straight, pror- low and appears to be differentiated into siradiate primary and secondary ribs termi- asymmetric clavi that are less numerous than nating in ventrolateral tubercles. Low keel the ribs. Ribs are narrow and prorsiradiate; differentiated into low asymmetric clavi they number 35 on the outer whorl, where slightly more numerous than the ribs. Inner one or two secondaries separate primaries. whorls usually densely ribbed. Primary ribs In the lower, Hoplitoides sandovalensis

← Fig. 70. Prionocyclus macombi Meek, 1876. A±C. USNM 498344; D, E. USNM 498394; F±H, USNM 498349, all gracile forms from locality 39. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_86 Allen Press • DTPro System File # 01235TQ

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Fig. 72. Prionocyclus macombi Meek, 1876. A, B. USNM 498397; C±E. USNM 498340; F, G. USNM 498398; H, I. USNM 498341, all robust forms from USGS Mesozoic locality 39. All ®gures are ϫ1.

Subzone of the Prionocyclus hyatti Zone, a In the robust form (®gs. 55C, D, 56±58), collection from a narrow horizon in the Sem- nuclei show great variation. Coiling is very illa Sandstone Member of the Mancos Shale evolute, with the umbilical seam indented to at USGS locality 28873 (locality 41 in ®g. accommodate the inner ventrolateral tuber- 1) is typical. It and other collections reveal cles of the inner whorl, the umbilicus is of that both robust and gracile forms are present moderate depth, the umbilical wall ¯attened in similar numbers. and subvertical, the umbilical shoulder quite

← Fig. 71. Prionocyclus macombi Meek, 1876. A, B. USNM 498348; C, D. USNM 498395, both gracile forms from locality 39. E, F. USNM 498396, robust form from locality 55. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_88 Allen Press • DTPro System File # 01235TQ

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narrowly rounded, the whorls slightly compressed to depressed, quadrate in intercostal section, and octagonal in costal section (table 7). At one extreme there are as few as 20 ribs per whorl, with up to 30 at the other. The ribs are all bullate primaries of equal development, strong, straight, and prorsiradiate, and strengthen across the ¯anks to strong conical to clavate inner ventrolateral Fig. 74. External suture of Prionocyclus ma- tubercles. A broad prorsiradiate rib connects combi Meek, 1876, from USNM 498854, from an inner ventrolateral tubercle to a strong locality 36. outer ventrolateral clavus, and a smooth zone separates the ribs from a strong siphonal keel, with clavi occupying the interspaces be- ribs are ¯ared, usually bullate, with inner and tween tubercles (®g. 56C±D). As size in- outer ventrolateral tubercles; the weaker ribs creases, the ribs become very coarse and lack bullae and have feeble or no inner ven- massive, with strong (®g. 56H±J) to weak trolateral tubercles. This style of ornament bullae (®g. 56C, D), with as few as 16 ribs may persist into middle growth, and there are per whorl. Sometimes the bullae are of var- gracile forms with dense, even ribs that may iable strength (®g. 56E±G), and displaced out reduce to mere striae (®g. 51N) and others from the umbilical shoulder. The outer ven- with ribbing of variable strength and tubercle trolateral clavi are assimilated into the inner development (®g. 52K, L). Inner and outer ventrolateral tubercles, which enlarge into ventrolateral tubercles remain separate and massive blunt horns that project outward but do not merge into horns, as in the robust usually not signi®cantly above the level of form; nonbullate ribs may lack an inner ven- the venter (®g. 56E±J). Other robust individ- trolateral tubercle and have much weaker uals have nuclei in which strong, bullate pri- outer ventrolateral tubercles than on the bul- mary ribs are separated by up to three weaker late primaries. The keel is broadly and irreg- nonbullate primaries, with a total of up to 45 ularly undulose. ribs per whorl. This differentiation persists Both robust and gracile forms show enor- into middle growth. The bullate primaries mous variation in ribbing and tuberculation have strong umbilical bullae and inner and style as well as timing of ontogenetic chang- outer ventrolateral tubercles. The nonbullate es in ornament. This is more readily appre- primaries sometimes connect to the umbilical ciated from the illustrations (®gs. 50±60) bullae as mere striae and have weak ventro- than from mere description. lateral tubercles, in some cases, only outer Prionocyclus hyatti from the upper part of ones. In some specimens (®g. 58) the stron- its zonal range is illustrated by collections ger primaries develop massive horns or from the Carlile Shale of Kansas, where the spines, with hornless ribs between. The larg- species occurs as fragments, often of large est robust individual at hand is 270 mm in body chambers, in ironstone concretions. diameter. Original aragonite shell is often preserved Nuclei of the gracile form (®gs. 51A±R, (®g. 57). Many of the specimens we have V±Y, 52A±L, 53, 54, 55E±J) overlap to a de- seen from Kansas are large, up to 300 mm in gree with those of the robust form. They are diameter. Juveniles are as variable as those more compressed, with a broad, shallow um- from the Hoplitoides sandovalensis Subzone bilicus (table 7), and have up to 50 ribs per in New Mexico and elsewhere already de- whorl that may be even (®g. 51M±O) or vary scribed, with even, coarse to ®ne ribbing as greatly in strength (®g. 51A±I). The stronger well as markedly differentiated weak and

← Fig. 73. Prionocyclus macombi Meek, 1876. A, B. USNM 498399; C, D. USNM 498342, both robust forms from USGS Mesozoic locality 39. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_90 Allen Press • DTPro System File # 01235TQ

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TABLE 8 Dimensions of juvenile Prionocyclus macombi Meek, 1876a

strong ribs. This variation is maintained into elements; E/L is broad and bi®d, and L is

middle growth, and most individuals have narrower and bi®d, as is L/U2. strong, bullate primary ribs with massive ven- DISCUSSION: Prionocyclus hyatti is the spe- trolateral horns and weaker, nonbullate inter- cies of the genus closest to Collignoniceras, calated ribs between that have weak ventro- discussed above. Juvenile stages show certain lateral tubercles. At the largest diameters, common characters, but middle and late fragments may bear as few as six ribs with growth stages are distinct. Prionocyclus hyatti massive ventrolateral horns per half whorl, differs from the most closely comparable and some specimens show the horns to have Prionocyclus, P. macombi, as follows: the ro- been septate, with a long spine (®g. 59). bust forms of the two species may have a de- These large specimens commonly show an gree of overlap in their coiling, whorl section, outward migration of the umbilical bullae, and expansion rate, but small and medium- which become conical and occupy an inner sized individuals of P. macombi never devel- ¯ank position. The keel of these specimens op ventrolateral horns as massive as those of is variable, from broadly undulose with P. hyatti, nor the striking differentiation into coarse clavi only slightly more numerous strong horned ribs with umbilical bullae, and than ventrolateral tubercles (®g. 57A, B) to weak, nonbullate hornless ribs. The gracile others where the crenulations are minute and forms of P. macombi are much more com- far more numerous. pressed, higher-whorled, and more involute The suture (®g. 60) has moderately incised than the corresponding forms of P. hyatti. The

TABLE 9 Dimensions of Prionocyclus macombi Meek, 1876a BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_91 Allen Press • DTPro System File # 01235TQ

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Fig. 75. Distribution of Prionocyclus macombi Meek, 1876, in the Western Interior Seaway during the middle Turonian. Land areas indicated by stipple. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_92 Allen Press • DTPro System File # 01235TQ

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Fig. 76. Prionocyclus bosquensis Kennedy, 1988. A. Paratype OUM KT 3858. B. Paratype OUM KT 3281. C. Paratype OUM KT 3801b. D. Holotype, OUM KT 3854. E. Paratype OUM KT 3869. All specimens are from the upper Turonian Scaphites whit®eldi Zone, South Bosque Formation, 10 km (6 mi) southwest of Waco, McLennan County, Texas. All ®gures are ϫ2.

ribbing is also denser and ®ner in P. macombi Western Interior from northern Montana with the ribs often reduced to mere striae, and southward into Trans-Pecos Texas and east- the inner and outer ventrolateral tubercles of ward from there into central and northern the inner whorls replaced by a single ventro- Texas (®g. 61). Also known from northern lateral clavus at an early stage. Chihuahua, Mexico, and Kazakhstan (Mar- OCCURRENCE: Widely distributed in the cinowski et al., 1996). BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_93 Allen Press • DTPro System File # 01235TQ

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Fig. 77. Prionocyclus bosquensis Kennedy, 1988. A. Paratype OUM KT 3870. B. Paratype OUM KT 3868. C. Paratype OUM KT 3873. All specimens are from the lower upper Turonian Scaphites whit®eldi Zone, South Bosque Formation, 10 km (6 mi) southwest of Waco, McLennan County, Texas. All ®gures are ϫ1.

Prionocyclus albinus (Fritsch, 1872) with ¯attened ¯anks. Ornament of numerous Figures 62, 108C prorsiradiate ribs with primaries usually separated by two or three secondaries. Ammonites albinus Fritsch, 1872: 28, pl. 6, ®g. 4. DESCRIPTION: The lectotype is 41 mm in Ammonites albinus Fritsch. Diener, 1925: 23. Prionocyclus hyatti (Stanton, 1894). Kennedy, 1988: diameter. Coiling is evolute, with U/D ϭ 75 (pars), pl. 15, ®gs. 3±5; text-®g. 25H±L only. 0.42. The umbilicus is shallow, with a feebly convex wall, and more narrowly rounded TYPE: Lectotype, here designated, is the umbilical shoulder. On the last half whorl, original of Fritsch, 1872: plate 6, ®gure 4, six or seven primary ribs arise on the um- no. 03144 in the Collections of the Narodni bilical wall, and strengthen into feeble bullae Museum, Prague, from the Turonian of Weh- perched on the umbilical shoulder. The ribs lovice, in the Czech Republic (®g. 108C is a are coarse, straight, and prorsiradiate on the plaster cast taken from an external mold of ¯anks, with inner and outer ventrolateral cla- the type). vi; the venter is not preserved. These strong DIAGNOSIS: A moderate-sized, fairly evo- primaries are separated by single much lute species that has whorls higher than wide weaker ribs that lack umbilical bullae and in- BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_94 Allen Press • DTPro System File # 01235TQ

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Fig. 78. Prionocyclus wyomingensis Meek, 1876. Lectotype, USNM 7729, middle Turonian Sca- phites warreni Zone, Wall Creek Member of the Frontier Formation near Medicine Bow, Carbon County, Wyoming. Figures are ϫ1.

ner ventrolateral tubercles. The poorly pre- four and as few as one of these weaker ribs served inner whorls are much more ®nely may separate the stronger primaries. Ribbing ribbed. of this type extends onto the body chamber Western Interior specimens referred to the of USNM 498339 (®g. 62P), interpreted as species have ®nely and evenly ribbed inner an adult. whorls (®g. 62A, D, F, L, O); the ribs are The suture is moderately incised, as in P. equal, crowded, straight, and prorsiradiate on hyatti. the ¯anks. This stage is succeeded, at vari- DISCUSSION: Prionocyclus albinus may able diameters, by a growth stage in which represent a paedomorphic derivative of the ribs differentiate into stronger primaries Prionocyclus hyatti in which the crowded or- with umbilical bullae, strong inner, and nament and variable rib strength of juveniles weaker outer ventrolateral tubercles, separat- of the latter are retained to maturity. Small ed by much weaker ribs that lack bullae and specimens of the two species are liable to be inner ventrolateral tubercles, but have feeble confused, but in middle growth, the change or effaced outer ventrolaterals; as many as to more distant ribbing in P. hyatti is dis- BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_95 Allen Press • DTPro System File # 01235TQ

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1988: 83, 84, pl. 2, ®g. 11; pl. 18, ®gs. 4±9; text-®gs. 28, 29b, e, j. Prionocyclus macombi Meek. Emerson et al., 1994: 212, 378. Prionocyclus macombi Meek, 1876. Reyment and Kennedy, 2001: ®gs. 2e±h, 4.

TYPES: The lectotype here designated is USNM 20259, the original of Meek, 1876b: plate 2, ®gure 3 (®g. 63A, B), from the Juana Lopez Member of the Carlile Shale of Colfax County, New Mexico, middle Turonian Prionocyclus macombi Zone. Meek refers to more than one specimen of Prionocyclus? macombi in his original account, indicating the species to be based on a type series. The lectotype is the original of his plate 2, ®gure 3a±c, and the original of his ®gure 3d may conceivably be the inner whorls of this spec- Fig. 79. Prionocyclus wyomingensis Meek, imen. 1876. Paralectotype, USNM 7729, middle Turon- DIAGNOSIS: A moderate-size species that ian Scaphites warreni Zone, Wall Creek Member occurs as an early form with arched venter of the Frontier Formation near Medicine Bow, and a later form with ¯attened venter. Keel Carbon County, Wyoming. Figures are ϫ1. low and notched into as many as three or four clavi per rib. The gracile form is excep- tinctive, and the two species reach maturity tionally compressed and has densely ribbed at widely differing diameters. innermost whorls; the outer adult whorls are OCCURRENCE: Prionocyclus hyatti Zone, nearly smooth. The robust form has coarser Smith County, Kansas; Emery County, Utah; and stronger ornament than the gracile form; Sandoval County, New Mexico; Dallas ribs support nodate to bullate umbilical tu- County, Texas. The type material is from the bercles and nodate to spinose inner ventro- Turonian of the Czech Republic. lateral tubercles, whereas outer ventrolateral clavi are barely perceptible or absent. Prionocyclus macombi Meek, 1876 DESCRIPTION: Robust and gracile forms of Figures 63±74 Prionocyclus macombi are distinct from the early growth stages on. During middle and Prionocyclus? macombi Meek, 1876b: 132, pl. 2, later growth stages, it is also possible to rec- ®g. 3a±d. ognize an early form of the species, where Prionocyclus? macombi Meek. Stanton, 1894: the venter is distinctively arched and rounded 172, pl. 41, ®gs. 1±5. Prionocyclus? macombi Meek. Logan, 1898: 264. (®gs. 68L, 71A±D, 72, 73) and a late form Prionocyclus macombi Meek. Johnson, 1903: in which the venter is ¯attened (e.g., ®g. 139. 68P). Prionocyclus macombi Meek. Grabau and Shimer, The earliest growth stages are highly var- 1910: 288, ®g. 1510e±g. iable and individuals may be nearly smooth Prionocyclus macombi Meek. Diener, 1925: 155. (®g. 67Y, Z) to densely ribbed (®g. 67A±E) Prionocyclus macombi Meek. Kauffman, 1977: in the gracile form. The smoothest variants pl. 26, ®gs. 4, 5. have distant umbilical bullae and tiny outer Prionocyclus macombi Meek. Hook and Cobban, ventrolateral clavi, even though ¯ank orna- 1980: 46, text-®g. 5A±G. ment is virtually absent. Densely ribbed ju- Prionocyclus macombi Meek. Hill, 1982: ®g. 22H, I. veniles generally have up to 16 strong pri- Prionocyclus macombi Meek. Cobban, 1984a: 86. mary ribs, sometimes feebly bullate, and giv- Prionocyclus macombi Meek. Cobban, 1986: ®g. ing rise to bundles of ribs. One to four weak- 7D, E. er primary and intercalated ribs occur Prionocyclus macombi Meek, 1876b. Kennedy, between the stronger ribs and extend to the BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_96 Allen Press • DTPro System File # 01235TQ

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Fig. 80. Prionocyclus wyomingensis Meek, 1876. Paralectotype, USNM 7729, middle Turonian Sca- phites warreni Zone, Wall Creek Member of the Frontier Formation near Medicine Bow, Carbon County, Wyoming. Figures are ϫ1.

umbilicus or arise on the inner ¯ank. Ribs whorl of three nuclei are given in table 8. At are prorsiradiate and straight to feebly ¯ex- this stage, the shell is compressed and high- uous across the ¯anks. They project forward whorled with a small umbilicus comprising and are concave over the ventrolateral shoul- as little as 17% of the shell diameter with der. Weak inner ventrolateral and clavate out- whorl breadth to height ratios of as little as er ventrolateral tubercles are present on the 0.5. The umbilical wall inclines outward and stronger primary ribs; the weaker primary merges with a broadly rounded umbilical and intercalated ribs are feebly or nontuber- shoulder. The ¯anks are ¯attened and sub- culate. This densely ribbed stage terminates parallel, and the ventrolateral shoulders are abruptly in most specimens by about 15±20 broadly rounded. The venter is arched in ear- mm diameter or less. ly forms, and more ¯attened in later ones. Dimensions, ratios, and number of ribs per Delicate umbilical bullae of variable strength

→ Fig. 81. Prionocyclus wyomingensis Meek, 1876. A, B. USNM 498409; C, D. USNM 498410, both gracile forms from locality 29. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_97 Allen Press • DTPro System File # 01235TQ

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Fig. 82. Prionocyclus wyomingensis Meek, 1876. USNM 7729D. This specimen bears the label 7729 in the same hand as the ®gured syntypes, and appears to be a previously unrecognized syntype, now paralectotype of the species. Middle Turonian Scaphites warreni Zone, Wall Creek Member of the Frontier Formation near Medicine Bow, Carbon County, Wyoming. Figure is ϫ0.9. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_99 Allen Press • DTPro System File # 01235TQ

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and generally 14±20 per whorl perch on the umbilical shoulder (®g. 67Q±X). They give rise to narrow, ¯exuous prorsiradiate ribs that terminate in small clavate to bullate inner ventrolateral tubercles. Outer ventrolateral tubercles may also be present, linked to the inner by prorsiradiate ribs, but disappear in early middle growth. The wide interspaces between these bullate primaries bear weak ribs that arise at the umbilical seam or low on the ¯anks plus lirae and growth striae (®g. 67Q±X). These weaker ribs may bear inner, but not outer ventrolateral tubercles, giving the shell a distinctive appearance when viewed ventrally (®g. 69B, C). Ornament sweeps forward and is concave over the ventrolateral shoulder, forming an acute ventral chevron. The keel is minutely and evenly serrated, the serrations far exceeding the ribs in number (®gs. 69A±C, F, G; 70A±C). The largest gracile forms are still septate at 150 mm diameter (table 9). These specimens are ornamented by distant, narrow prorsiradiate ribs with weak umbilical bullae, up to 26 per whorl, separated by broad interspaces that are smooth on molds. Most specimens only have inner ventrolateral clavi at this stage, while there are some specimens with clavi intercalated between the ribs or even short intercalated ribs. Riblets and striae form an acute chevron on the venter, while the keel remains minutely serrated when well preserved. Robust forms have a much wider umbilicus and the whorl section varies from slightly compressed to depressed. In juveniles (®g. 67F±P) there are usually approximately 12 prominent umbilical bullae per whorl. These give rise to distant, narrow, rounded, straight, prorsiradiate primary ribs separated by wide interspaces, smooth on most molds. Bullae may be equal in strength or vary, while a few specimens have shorter intercalated ribs (®g.

← 1876. USNM 7729D. This specimen bears the label 7729 in the same hand as the ®gured syntypes, and appears to be a previously unrecognized syntype, now paralectotype of the species. Middle Turonian Scaphites warreni Zone, Wall Creek Member of the Frontier Formation near Medicine Fig. 83. Prionocyclus wyomingensis Meek, Bow, Carbon County, Wyoming. Figure is ϫ0.9. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_100 Allen Press • DTPro System File # 01235TQ

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Fig. 84. Prionocyclus wyomingensis Meek, 1876. A, B. USNM 498411, robust form from locality 29. C, D. USNM 356921, robust form from locality 37. E, F. USNM 498412, robust form from locality 33. G, H. USNM 498413; I, J. USNM 498414, both gracile forms from locality 29. All ®gures are ϫ1.

69H, I). All ribs terminate in conical to fee- ventrolateral clavi, which are lost early in bly clavate inner ventrolateral tubercles, middle growth. The ventral keel is entire and which may be equal or of variable strength. minutely serrated. If inner ventrolateral tu- Broad ribs sweep forward to feeble outer bercles are strong, the umbilical seam and BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_101 Allen Press • DTPro System File # 01235TQ

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Fig. 85. Prionocyclus wyomingensis Meek, 1876. A±C. USNM 498403, gracile form from locality 28. D, E. USNM 498401, robust form from locality 28. F, G. USNM 498416, gracile form from locality 29. H, I. USNM 498402, robust form from locality 29. All ®gures are ϫ1.

wall of the succeeding whorl may be notched As size increases, ornament may coarsen to accommodate them (®g. 69H). The venter markedly, and a common variant has a de- is rounded or arched in early forms (®g. 68J± pressed, polygonal coastal whorl section, L), ¯attened in later ones (®g. 68F±I, O, P). strong conical umbilical bullae displaced out BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_102 Allen Press • DTPro System File # 01235TQ

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Fig. 86. Prionocyclus wyomingensis Meek, 1876. Pathological specimen lacking keel. USNM 1 475903, ex J.I. Kirkland Collection, from the Juana Lopez Member of the Mancos Shale, in the SE /4 sec. 4, T. 12S, R. 1E, Gar®eld County, Colorado. Figures are ϫ1.

Fig. 87. External suture line of Prionocyclus wyomingensis Meek, 1876, USNM 498411, from locality 29. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_103 Allen Press • DTPro System File # 01235TQ

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TABLE 10 Dimensions of Prionocyclus wyomingensis Meek, 1876a

a little from the umbilical wall, low, broad, lae and a sharpening of ribs plus strong distant radial ribs, effaced at mid¯ank, and growth striae and lirae on the umbilical wall blunt, conical inner ventrolateral horns. suggesting maturity at a size of 220±250 There may be low, blunt ventrolateral ribs mm diameter. The fairly simple suture (®g. with feeble tubercles between (®gs. 72H, I). 74) has been illustrated by Meek (1876b: pl. The keel commonly lacks regular serrations 2, ®g. 3c) and by Stanton (1894: pl. 41, ®g. on the internal mold. 3). The largest fragments of the robust form DISCUSSION: Differences between Priono- are still septate at whorl heights of 70 mm, cyclus macombi and P. hyatti are discussed corresponding to an estimated diameter of above. Prionocyclus bosquensis Kennedy, 220 mm (table 9). Ornament consists of 1988 (86, pl. 18, ®gs. 1±3, 10, 11; text-®g. coarse, distant bituberculate ribs, the inner 29a, d, f±i; ®gs. 76, 77), which is known only ventrolateral tubercles developing into sharp from crushed specimens, differs from P. ma- horns on molds, while there are occasional combi in showing bunching of ribs at um- feeble intercalated ventrolateral tubercles. A bilical bullae as well as persistence of inner body chamber fragment with a whorl height and outer ventrolateral tubercles in middle of 90 mm shows weakening umbilical bul- growth. P. wyomingensis (®gs. 78±87) is

TABLE 11 Dimensions of Prionocyclus novimexicanus (Marcou, 1858)a BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_104 Allen Press • DTPro System File # 01235TQ

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Fig. 89. Prionocyclus novimexicanus (Marcou, 1858). A, B. Holotype of Prionocyclus wyomingensis elegans Haas, 1946, the original of Whit®eld, 1880, plate 14, ®gures 1, 2. C, D. Original of Whit®eld, 1880, plate 14, ®gure 3. Both specimens are registered as USNM 12283, ``in silico-calcareous layers of Division No. 2, Cretaceous, on the east fork of Beaver Creek, near Camp Jenny, Black Hills.'' All ®gures are ϫ1.

known mostly from the gracile form, which Prionocyclus bosquensis Kennedy, 1988 exhibits much stronger, coarser ribbing than Figures 76, 77 that of P. macombi with slender whorls. In Prionocyclus bosquensis Kennedy, 1988: 86, pl. addition, in P. wyomingensis the ribs some- 18, ®gs. 1±3, 10, 11; text-®gs. 29a, d, f±i. times arise in groups from bullae, and an in- Prionocyclus bosquensis Kennedy, 1988. Emer- ner lateral bulge or tubercle and a ®ngerlike son et al., 1994: 212, 378. septate horn on the inner ventrolateral tuber- TYPES: Holotype is OUM KT 3854 (®g. cle develop at maturity. 76D); there are numerous paratypes of which OCCURRENCE: Prionocyclus macombi has the following are illustrated: OUM KT 3281 been found in a wide belt paralleling the (®g. 76B), 3801b (®g. 76C), 3858 (®g. 76A), western shoreline of the basal late Turonian 3868 (®g. 77B), 3869 (®g. 76E), 3870 (®g. seaway that extends from western South Da- 77A), and 3873 (®g. 77C), from the top 6±7 kota southwestward to central Utah and then m of the South Bosque Formation, Cement southeastward through most of New Mexico Works, 10 km (6 mi) southwest of Waco, (®g. 75). The species is also known from McLennan County; also OUM KT 5375±85, north-central Texas. pyritic juveniles from the same locality and

← Fig. 88. Prionocyclus novimexicanus (Marcou, 1858). A±C. BMNH C49764, lectotype, a gracile form from either near Albuquerque, New Mexico, or La Lunes, New Mexico. D. USNM 498424, gracile form from locality 2. E, F. USNM 498421, gracile form from locality 1. G. USNM 498418, gracile form from locality 2. H, I. USNM 498425, gracile form from locality 1. J. USNM 498426, gracile form from locality 2. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_106 Allen Press • DTPro System File # 01235TQ

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Fig. 91. Prionocyclus novimexicanus (Marcou, 1858). A, B. USNM 254556, gracile form from locality 13. C, D. USNM 498431, gracile form from locality 2. All ®gures are ϫ1.

horizon; OUM KT 5372a, b collected in situ minutely serrated siphonal keel. A few spec- 3 m below the base of the Austin Chalk at imens may even lack differentiated ribs and this locality; OUM KT 5373a, b from 1 m bear ®ne, even ornament, without tubercles; below the Austin Chalk; upper Turonian Sca- some individuals show groups of 3 or so ribs phites whit®eldi zone. arising from a bulla while all specimens bear DIAGNOSIS: A small densely ribbed species long intercalated ribs inserted on the inner with a minutely notched keel. Ribs ¯exuous ¯ank. These specimens usually retain their and not distinctly separated into primaries outer ventrolateral tubercles. They are linked, and secondaries. by strengthening of tubercles and progressive DESCRIPTION: This species has been fully decline of the weaker, feebly tuberculate ribs described by Kennedy (1988: 86, 87): ``Most to much rarer individuals with distant ribs specimens are 10±45 mm diameter, are with umbilical bullae, strong inner and weak crushed, and retain original aragonitic shell. outer ventrolaterals, the interspaces with Coiling is fairly involute, with U ϭ 29%. Or- weak to obsolete ¯ank ribs, but short inter- nament is highly variable. At one extreme calated ribs on the ventrolateral shoulders are very ®nely ornamented individuals. and venter. There are a few much larger frag- These have very dense, crowded, narrow, ments, with whorl heights of up to 53 mm. ¯exuous prorsiradiate ribs, every ®fth to All of these show strong distant ribs with sixth strengthened, with a tiny umbilicolater- umbilical bullae, prominent inner ventrolat- al bulla, a bullate inner ventrolateral, and an eral spines or tubercles, weak, or no outer obliquely placed outer ventrolateral tubercle. ventrolaterals, and groups of riblets and The intermediate ribs generally lack umbili- growth striae extending from the inner ven- cal bullae, and have very faint or no inner trolateral nodes towards the mid-venter. Be- and outer ventrolaterals. There is a strong, tween these ribs are much ®ner intercalated

← Fig. 90. Prionocyclus novimexicanus (Marcou, 1858). A±C. USNM 498419; D, E. USNM 498427, both from locality 2. F, G. USNM 498420, gracile form from locality 3. H, I. USNM 498428, gracile form from locality 2. J, K. USNM 498429, gracile form from locality 3. L. USNM 498430, gracile form from locality 2. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_108 Allen Press • DTPro System File # 01235TQ

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Fig. 92. Prionocyclus novimexicanus (Marcou, 1858). A±C. USNM 498432; D, E. USNM 498433; F, G. USNM 254555, all gracile forms from locality 13. H±J. USNM 498434, a gracile form from locality 24. K, L. USNM 498436, a gracile form from locality 31. M, N. USNM 498435, a gracile form from locality 13. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_109 Allen Press • DTPro System File # 01235TQ

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TABLE 12 Dimensions of Prionocyclus germari (Reuss, 1845)a

riblets and lirae. There is a strong crenulate Prionocyclus wyomingensis Meek. Logan, 1898: siphonal keel. Sutures not seen.'' 463, pl. 106, ®gs. 1±4. DISCUSSION: See Kennedy (1988: 88). Prionocyclus wyomingensis Meek. Johnson, 1903: OCCURRENCE: As for types. 139. Prionocyclus wyomingensis Meek. Barbour, 1903: pl. 5, ®gs. 4, 5. Prionocyclus wyomingensis Meek, 1876 Prionocyclus wyomingensis Meek. Grabau and Figures 78±87 Shimer, 1910: 228, ®g. 1510a±d. Ammonites serrato-carinatus Meek, 1870: 429. Prionocyclus wyomingensis Meek. Diener, 1925: Ammonites (Pleuroceras?) serrato-carinatus 155. Meek, 1871: 298. Prionocyclus reesidei Sidwell, 1932: 318 (pars), Prionocyclus wyomingensis Meek, 1876a: 452, pl. 49, ®gs. 10, 11 only. footnote. Prionocyclus wyomingensis Meek. Roman, 1938: Prionocyclus wyomingensis Meek. White, 1880: 457, pl. 46, ®g. 435. 35, pl. 15, ®g. 1a±e. Prionocyclus wyomingensis Meek. Shimer and non Prionocyclus wyomingensis Meek. Whit®eld, Shrock, 1944: 593, pl. 247, ®gs. 3, 4. 1880: 440, pl. 14, ®gs. 1±3 (ϭ P. novimexican- Prionocyclus wyomingensis Meek. Haas, 1946 us (Marcou). and varieties robusta (p. 200), non var. elegans Prionocyclus wyomingensis Meek. White, 1883: (p. 210) (ϭ P. novimexicanus); pl. 18, ®gs. 3± 35, pl. 15, ®g. 1a±e. 6; non pl. 19, ®gs. 1±7, 11±14 (ϭ P. novimex- Prionocyclus wyomingensis Meek. Boyle, 1893: icanus); pl. 20, ®gs. 1±3, 5±7, non pl. 20, ®g. 243. 4(ϭ P. novimexicanus); pl. 22, ®gs. 3±5; non Prionocyclus wyomingensis Meek. Stanton, 1894: pl. 22, ®gs. 1, 2 (ϭ P. novimexicanus); pl. 23, 171, pl. 40, ®gs. 1±4. ®gs. 1, 3 (in part); pl. 24, ®gs. 2, 3; text-®gs. Prionocyclus wyomingensis Meek. Gilbert, 1896: 93±97, 105±108; non 98±104 (ϭ P. novimexi- 565, pl. 58, ®gs. 1±3. canus).

TABLE 13 Dimensions of Prionocyclites mite Kennedy, 1988a BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_110 Allen Press • DTPro System File # 01235TQ

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Prionocyclus wyomingensis Meek. Wright, 1957: vate to bullate outer ventrolateral tubercles. 426, ®g. 547. 6a, 6b. Robust form has coarse ribs that support Prionocyclus wyomingensis Meek. Termier and prominent nodate umbilical tubercles, nodate Termier, 1960: ®g. 2418. inner ventrolateral tubercles, and weak outer Prionocyclus wyomingensis Meek. Matsumoto, ventrolateral clavi. 1965: 18, pl. 16, ®g. 1a±c; pl. 17, ®g. 2a±c; pl. 18, ®g. 1a±c (? ϭ P. novimexicanus). DESCRIPTION: The robust form is rare (®gs. Prionocyclus wyomingensis Meek. Matsumoto, 84A±F, 85D, E, H, I). Coiling is evolute with 1971: 132, pl. 21, ®g. 2; pl. 22, ®g. 1, text-®g. U/D ϭ 0.30±0.37 in the available material 2. (table 10); the umbilicus broad, the umbilical Prionocyclus wyomingensis Meek. Hattin, 1975b: wall ¯attened and inclined outward, with pl. 2, ®g. 11. grooves to accommodate the inner ventrolat- Prionocyclus wyomingensis Meek. Kauffman, eral spines of the preceding whorl (®g. 85E). 1977: pl. 26, ®gs. 2, 3. The umbilical shoulder is narrowly rounded, Prionocyclus wyomingensis Meek, 1876. Kenne- the whorl section slightly compressed to dy et al., 1989: 91, ®g. 24E. slightly depressed, with the greatest breadth Prionocyclus wyomingensis Meek, 1876. Kenne- dy, 1988: 88, pl. 14, ®g. 8. at the umbilical bullae costally and just out- Prionocyclus wyomingensis Meek, 1876. Hall et side the umbilical shoulder intercostally. al., 1994: 305, ®gs. 4I, N, O, 5A±C, G±J. Ornament is highly variable. At one ex- Prionocyclus wyomingensis Meek. Emerson et al., treme are very coarsely ribbed individuals 1994: 212, 378. with only 16 or 17 ribs per whorl. Ribs arise Prionocyclus wyomingensis Meek. Wright, 1996: at the umbilical seam and strengthen into 186, ®g. 4a, b. massive conical umbilical bullae, broaden Prionocyclus wyomingensis Meek, 1876. Rey- and are straight and prorsiradiate across the ment and Kennedy, 2001: ®g. 3. ¯anks, and develop massive septate inner TYPES: Lectotype, by the subsequent des- ventrolateral spines that generally appear as ignation of Matsumoto, 1965: 18 is USNM large clavi on the mold when the spines have 7729, the original of Stanton, 1894: plate 40, broken off. The rib pro®le is somewhat con- ®gure 3 (®g. 78); two paralectotypes have the cave between bulla and spine (®g. 85H, I). same number and are the originals of Stan- Broad ribs sweep forward across the ventro- ton, 1894: plate 40, ®gures 1, 2 (®gs. 79, 80). lateral shoulder to outer ventrolateral clavi A further probable un®gured paralectotype and a smooth zone separates them from the bears the number 7729D, and is shown in strong and minutely serrated siphonal keel. ®gures 82, 83. All are from what is now At the other extreme (®g. 85D, E) are sim- known as the Wall Creek Sandstone Member ilarly proportioned individuals that have 17 of the Frontier Formation in the Medicine sharp to spinose umbilical tubercles in mid- Bow River valley of Southeast Wyoming dle growth, each of which gives rise to a (®de Cobban, 1984a: 86). strong concave to straight and rectiradiate DIAGNOSIS: A large species for the genus; adapical primary rib and a much weaker ada- moderately evolute; keel notched into more pertural primary rib, which loop to a strong siphonal clavi than ribs. Gracile form has inner ventrolateral clavus/spine, the arrange- closely spaced prorsiradiate primary and sec- ment resembling a bow. Broad ribs sweep ondary ribs. Primary ribs bear lengthy um- forward over the ventrolateral shoulder and bilical bullae that may divide into shorter connect to weaker outer ventrolateral clavi. umbilical bullae and bullate or nodate inner Between the looped, bullate ribs are one or lateral tubercles. All primaries bear nodate two nonbullate ribs that extend to the umbil- inner ventrolateral tubercles and weaker cla- ical seam or arise on the inner ¯anks and

← Fig. 93. Prionocyclus novimexicanus (Marcou, 1858). A, B. USNM 498437, a gracile form from locality 2. C, D. USNM 498438, a gracile form from locality 3. E, F. USNM 498439, a gracile form from locality 3. G. USNM 498440, a gracile form from locality 18. H. USNM 498441, a gracile form from locality 25. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_112 Allen Press • DTPro System File # 01235TQ

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Fig. 94. Prionocyclus novimexicanus (Marcou, 1858). USNM 498422, a gracile form from locality 2. Reduced ϫ0.53. Original is 386 mm in diameter. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_113 Allen Press • DTPro System File # 01235TQ

2001 KENNEDY ET AL.: COLLIGNONICERATINAE 113

Fig. 95. Prionocyclus novimexicanus (Marcou, 1858). USNM 498422; see also ®gure 94. Reduced ϫ0.53. Original is 386 mm in diameter. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_114 Allen Press • DTPro System File # 01235TQ

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Fig. 96. Prionocyclus novimexicanus (Marcou, 1858). USNM 498423, robust form from locality 4. Reduced ϫ0.53. The original is 380 mm in diameter. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_115 Allen Press • DTPro System File # 01235TQ

2001 KENNEDY ET AL.: COLLIGNONICERATINAE 115

develop much weaker ventrolateral tubercles, the arrangement giving the shell a highly irregular ornament. The largest robust individual seen is 83 mm in diameter. Gracile individuals are common (®gs. 78± 81; ?82, ?83; 84G±J, 85A±C, F, G). At 40 mm diameter, coiling is evolute, with U/D ϭ 0.26 (table 10); the umbilicus is shallow, with a low ¯attened umbilical wall. The whorls are high and compressed (whorl breadth to height ratio averages 0.79), with ®ne, dense, crowded ribs. There are up to 12 weak umbilical bullae that give rise to single or paired ribs, with single nonbullate ribs between that extend to the umbilicus or not, yielding a total of 44±48 ribs per whorl. Ribs are crowded, ¯exuous, and prorsiradiate, with small bullate to clavate inner, and clavate outer ventrolateral tubercles. In middle growth, specimens have up to 70 ribs per whorl. There are up to 18 bullae per whorl, initially at the umbilical shoulder, but tending to move out to the inner ¯anks, when a new bulla develops (®g. 81C). They are the bases of septate umbilical spines, and give rise to pairs of coarse ribs that are straight and prorsiradiate and initially loop to outer ventrolateral tubercles that are also the bases of septate spines. As size increases, the tendency to loop declines and the adapertural rib of the pair may detatch. There are one to six weaker, nonbullate long ribs between the bullate ones, each of which may develop a weak inner ventrolateral tubercle. The ribs sweep forward over the ventrolateral shoulder to small, obliquely placed outer ventrolateral tubercles and form an acute chevron with the high and minutely serrated siphonal keel. The largest gracile individuals are 220±230 mm diameter. At maturity, the bullate ribs strengthen markedly and are very coarse, high, and barlike. Where well-preserved, they bear long inner lateral and inner ventrolateral spines, while sharp outer ventrolateral clavi are also present. Be- tween are up to three much weaker, narrow, sharp long ribs that lack any trace of a tu-

← cou, 1858). USNM 498423, see also Figure 96. Reduced ϫ0.53. The original is 380 mm in di- Fig. 97. Prionocyclus novimexicanus (Mar- ameter. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_116 Allen Press • DTPro System File # 01235TQ

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bercle. This coarseness and marked differ- Haas, 1946: 200, pl. 19, ®gs. 1±7, 11±14; pl. entiation of the main ribs gives a highly char- 20, ®g. 4; pl. 21, ®gs. 1±3, 5; pl. 22, ®gs. 1, 2; acteristic appearance to the shell. text-®gs. 98±104. The largest collection studied consists of Prionocyclus wyomingensis Meek. Luppov and 73 specimens from the Frontier Formation at Druschchits, 1958: 123, text-®g. 97b, pl. 60, ®g. 3. USGS Mesozoic locality D9244 in south- Prionocyclus (Prionocyclus) wyomingensis var. eastern Wyoming. Of these specimens 61 are elegans Haas. Jeletzky, 1970: pl. 26, ®g. 8. the gracile form and 12 are the robust form. Prionocyclus wyomingensis Meek. Kennedy and The suture is fairly simple and typical of Cobban, 1976: pl. 11, ®g. 4. the genus. The suture from White (1880) is Prionocyclus wyomingensis elegans Haas. Kauff- shown here (®g. 87). man, 1977: 260, pl. 22, ®g. 14. DISCUSSION: Differences between P. Prionocyclus novimexicanus (Marcou). Hook and wyomingensis, P. macombi, and P. bosquen- Cobban, 1979: 38, ®g. 3E±L. sis are noted above. P. wyomingensis most Prionocyclus novimexicanus (Marcou). Mere- closely resembles P. novimexicanus (Mar- wether et al., 1979: pl. 3, ®g. 3. Prionocyclus wyomingensis Meek. Case, 1982: cou, 1858) (®gs. 88±101), which is presum- ®g. 12. 63. ably its descendant, but the latter is even Prionocyclus novimexicanus (Marcou). Cobban, more compressed, has ¯exuous ribs, seldom 1984b: 14, pl. 1, ®gs. 15, 16. develops the umbilical and inner lateral tu- Prionocyclus novimexicanus (Marcou, 1853). berculation of P. wyomingensis and loses the Kennedy, 1988: 83. outer ventrolateral tubercles at a very early Prionocyclus novimexicanus (Marcou, 1848). stage so that the primary ribs are effectively Kennedy et al., 1989: 93. bituberculate. Prionocyclus novimexicanus (Marcou). Cobban A remarkable pathological specimen (col- and Hook, 1989: ®g. 10j±l. lected by J. I. Kirkland, Salt Lake City, Prionocyclus novimexicanus (Marcou, 1858). AmeÂdro, 1990: 269, pl. 17, ®g. 6; pl. 25, ®gs. Utah), USNM 475903 (®g. 86), from the Ju- 1±3. ana Lopez member of the Mancos Shale in Prionocyclus novimexicanus (Marcou). Emerson the SE¼ sec. 4, T. 12 S, R. 1 E, Gar®eld et al., 1994: 212, 378. County, Colorado, lacks a siphonal keel (®g. Prionocyclus novimexicanus (Marcou, 1858). 86). Reyment and Kennedy, 2001: ®g 2i±m. OCCURRENCE: Prionocyclus wyomingensis is widely distributed in the Western Interior TYPES: The lectotype, here designated, of of the United States from northwestern Mon- Ammonites novimexicani Marcou, 1858, is tana south through much of New Mexico, BMNH C49764 (®g. 88A±C), either from and from east-central Utah to southwestern near Albuquerque, New Mexico, or La Kansas. It has also been ®gured recently Lunes, New Mexico. Marcou speci®cally from Alberta, Canada (Hall et al., 1994) and mentions two specimens in his account, and is known from Japan (Matsumoto, 1971). lectotype designation is necessary. The holotype of Prionocyclus wyomingensis ele- Prionocyclus novimexicanus (Marcou, gans Haas, 1946 (a synonym), is the original 1858) of White®eld, 1880: plate 14, ®gure 2, from Figures 88±101 Beaver Creek, near Camp Jenney, South Da- Ammonites novi-mexicani Marcou, 1858: 35, pl. kota, USNM 12283 (®g. 89A, B). 1, ®g. 2. DIAGNOSIS: This large, moderately evolute Prionocyclus wyomingensis Meek. Whit®eld, species has a keel notched into small siphon- 1880: 440, pl. 14, ®gs. 1±3. al tubercles more numerous than the ribs. Prionocyclus wyomingensis Meek var. elegans The gracile form has dense, ¯exuous primary

← Fig. 98. Prionocyclus novimexicanus (Marcou, 1858). A. USNM 498442, a gracile form from locality 2. B, C. USNM 498443, a gracile form from locality 9. D, E. USNM 498444, a robust form from locality 10. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_118 Allen Press • DTPro System File # 01235TQ

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and secondary ribs. Primary ribs arise from breadth at the umbilical bullae in costal sec- elongated umbilical bullae, each of which is tion and just outside the umbilical shoulder rarely divided into a shorter umbilical bulla intercostally. and a small, nodate inner lateral tubercle. All The earliest growth stages seen are char- primaries bear rounded to clavate inner ven- acterized by very ®ne, dense, crowded or- trolateral tubercles. Outer ventrolateral tuber- nament. Feeble umbilical bullae give rise to cles are usually absent, and very weak if pre- pairs or groups of three ribs; between are up sent. The robust form is more evolute and to nine ribs which arise at the umbilical more sparsely ribbed. Looped ribs may con- shoulder, low, or high on the ¯anks. The de- nect umbilical and inner ventrolateral tuber- gree of differentiation between strong and cles in both gracile and robust forms. bullate and weak and nonbullate ribs varies DESCRIPTION: The best example of the ro- greatly between individuals (®gs. 90±92), bust form is USNM 498417 (®g. 99F, G). with up to 90 ribs per whorl. In middle Coiling is very evolute, with U/D ϭ 0.48, growth, to a diameter of 70±80 mm, ribbing the umbilical wall ¯attened and inclined out- varies greatly in strength and density be- ward, the umbilical shoulder broadly round- tween individuals (compare ®g. 90L and ®g. ed, the ¯anks convergent with the greatest 99A, B). Typically, there are 14±18 bullae breadth at the umbilical shoulder intercostal- per whorl that give rise to pairs or groups of ly and at the umbilical bulla in costal section. ribs, with nonbullate intercalaries between, The whorl section is compressed, with a cos- the bullate ribs looping in pairs to inner ven- tal whorl breadth to height ratio of 0.73 up trolateral clavi, the nonbullate ribs with fee- to a shell diameter of 60 mm (table 11). Or- ble inner ventrolateral tubercles or not. Fee- nament consists of blunt umbilical bullae, 18 ble outer ventrolateral tubercles are present per whorl. These give rise to broad, low, only at diameters of less than 15±20 mm. straight prorsiradiate ribs that weaken mark- Beyond, ribs sweep forward over the ventro- edly at mid¯ank, and connect to blunt inner lateral shoulder to form an acute chevron. ventrolateral bullae. Broad ribs sweep for- The siphonal keel is high, strong, and mi- ward and decline, forming an acute chevron nutely serrated. with the minutely serrated siphonal keel; An adult of the gracile form has a maxi- there is no trace of an outer ventrolateral tu- mum diameter of 386 mm (®gs. 94, 95). It bercle. Beyond 60 mm, the bullae weaken retains the crowded ornament of ribs of vary- progressively, and commonly give rise to ing strengths to a diameter of 170 mm, be- pairs of ribs, which may loop to the inner yond which the minor ribs decline progres- ventrolateral bullae (®g. 100C, D) or not. Be- sively so that by the middle section of the tween are one or two weaker nonbullate ribs, body chamber, ornament consists of very dis- arising at the umbilical shoulder or low on tant, coarse, convex prorsiradiate ribs with the ¯anks with feeble inner ventrolateral tu- strong, subspinose umbilical bullae and enor- bercles or not. The ribs become feebly ¯ex- mous, slender septate inner ventrolateral uous at this stage. horns that persist to the last section of the The gracile form is represented by numer- body chamber. Just before the adult aperture, ous specimens (®gs. 88±95, 98A±C). Coiling ribbing stops and the last section is covered is moderately evolute, with a shallow umbi- in dense irregular riblets, lirae, and striae. licus with U/D ϭ 0.26±0.30 between 50 and USNM 498423 (®gs. 96, 97) is an un- 100 mm shell diameter; the umbilicus is shal- crushed body chamber mold. The whorl low, with a ¯attened wall. The whorl section height is 98 mm at the end of the phragmo- is compressed, with whorl breadth to height cone, and the diameter of the specimen is ratios of 0.5±0.7 (table 11), the greatest 380 mm, which suggests a total diameter of

← Fig. 99. Prionocyclus novimexicanus (Marcou, 1858). A, B. USNM 498445, a robust form from locality 2. C±E. USNM 498446, a robust form from locality 30. F, G. USNM 498417, a robust form from locality 1. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_120 Allen Press • DTPro System File # 01235TQ

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Fig. 100. Prionocyclus novimexicanus (Marcou, 1858). A, B. USNM 498447, a robust form from locality 2. C, D. USNM 498448, a robust form from locality 3. E±G. USNM 498449; H, I. USNM 498450, both robust forms from locality 31. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_121 Allen Press • DTPro System File # 01235TQ

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Fig. 101. External suture of Prionocyclus novimexicanus (Marcou, 1858). Copy of Haas, 1946, ®gure 104.

Fig. 102. Prionocyclus quadratus Cobban, over 400 mm. The whorl section at the end 1953. Paratype USNM 108334, Sage Breaks Mem- of the phragmocone is trapezoidal with a ber of the Carlile Shale, 14.9 km (9 mi) south-south- whorl breadth to height ratio of 0.95. Orna- east of Rapid City, in the NE¼ sec. 22, T. 1S, R. ment consists of strong, distant, coarse pri- 8W, Pennington County, South Dakota. Figure is mary ribs, with umbilical bullae that decline ϫ1. in strength as size increases. There are massive conical inner ventrolateral tubercles at over much of Colorado and New Mexico. It the end of the phragmocone and beginning has also been recorded from Trans-Pecos of the body chamber. They weaken on the Texas (Kennedy et al., 1989), and Tunisia later part of the body chamber. Narrow ribs (AmeÂdro, 1990). project forward over the venter to form a chevron, with ®ner and irregular riblets be- Prionocyclus quadratus Cobban, 1953 tween. The siphonal keel is blunt on the Figures 102±107 mold, and irregularly notched. The suture has a broad, bi®d E/L, narrow Prionocyclus reesidei Sidwell, 1932: 354 (pars.), L, and bi®d L/U (®g. 101). pl. 49, ®g. 12 only. 2 Prionocyclus quadratus Cobban, 1953: 354, pl. DISCUSSION: Prionocyclus novimexicanus 48, ®gs. 1±8. most closely resembles its presumed ances- Prionocyclus reesidei Sidwell. Kauffman, 1977: tor, P. wyomingensis (®gs. 78±87); differenc- 260, pl. 2, ®gs. 2, 3. es are discussed above. Prionocyclus qua- Prionocyclus quadratus Cobban. Kauffman, dratus (®gs. 102±107) and P. germari (®gs. 1977: 256, pl. 19, ®gs. 11, 12; pl. 26, ®g. 1. 109±119) never show the marked differen- non Prionocyclus quadratus Cobban. Merewether tiation of rib strength of juvenile P. novi- et al., 1979: pl. 3, ®gs. 6±8 (ϭ P. germari). mexicanus; instead the ribbing is even. The Prionocyclus quadratus Cobban, 1953. Kennedy, whorls are never as depressed or as ®nely 1988: 83. ribbed, although P. quadratus and P. ger- Prionocyclus quadratus Cobban. Cobban and Hook, 1989, ®g. 10H, I. mari lose their outer ventrolateral clavi at very small diameters, as does P. novimexi- TYPES: Holotype is USNM 108332 (®g. canus. Prionocyclus quadratus develops a 103), paratypes USNM 108333a±c (®gs. pronounced outer lateral bulge when adult 104±106), from the upper Turonian Sage (®gs. 116±118), while the umbilical bullae Breaks Member of the Carlile Shale in sec. commonly migrate out to an inner ¯ank po- 33, T. 9 S, R. 61 E, Carter County, Mon- sition in middle and late growth. tana. Paratype USNM 108334 (®g. 102) is OCCURRENCE: Widely distributed in the from a septarian concretion bed 12.8 m Western Interior in the upper Turonian Zone above the base of the Sage Breaks Member of Scaphites whit®eldi. Specimens have been 14.5 km south-southeast of Rapid City in found in south-central and southeastern the NE¼ sec. 22, T. 1 S, R. 8 W, Pennington Montana, central and eastern Wyoming, County, South Dakota. Paratype USNM western South Dakota, eastern Utah, and 108335 is from a septarian concretion bed BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_122 Allen Press • DTPro System File # 01235TQ

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Fig. 103. Prionocyclus quadratus Cobban, 1953. Holotype, USNM 108332, Sage Breaks Member of the Carlile Shale, sec. 63, T 9S, R 61E, Carter County, Montana. Figures are ϫ1.

5.2 m below the top of the Turner Sandy bing consists of strong, narrow, prorsiradi- Member of the Carlile Shale 10.4 km north ate primaries alternating with very weak, of Belle Fourche, in the E½ sec. 2, T. 9 N, narrow secondaries. All ribs are straight on R. 2 E, Butte County, South Dakota. crossing the ¯anks, but they bend sharply DIAGNOSIS: A fairly large species that has forward on the ventrolateral shoulder and sparsely ribbed adult whorls with strong pri- fade out on the venter. Primary ribs support mary ribs and weak secondary ribs. Primary umbilical bullae, weak mid¯ank bullae, and ribs on the robust form have umbilical bul- blunt ventrolateral tubercles. Secondary ribs lae, nodate inner ventrolateral tubercles, and arise low on the ¯anks and efface on the mid¯ank swellings or weak tubercles. The venter without developing into ventrolateral gracile form usually lacks mid¯ank swell- tubercles. ings. No clearly de®ned outer ventrolateral A paratype, USNM 108334 (®g. 102), tubercles occur on either form. about 43 mm in diameter, has 42 ribs per DESCRIPTION: The types represent the ro- whorl at a diameter of 30 mm with two or bust form of the species. The holotype, three secondaries separating primaries. At a USNM 108332 (®g. 103), is part of a small larger diameter, most primaries are separated adult body chamber that has an intercostal by a single secondary. width of 32.5 mm and height of 34.3 mm The suture ®gured by Cobban (1953: pl. (ratio ϭ 0.95) and a costal width of 42.0 mm 48, ®g. 4) has a rather long lateral lobe, and and height of 38.2 mm (ratio ϭ 1.10), with a similar feature is shown in ®gure 107. the greatest width at the umbilical shoulder. DISCUSSION: The species attains a large The specimen is an internal mold that does size, with some phragmocones having di- not have serrations on the keel preserved. A ameters of as much as 250 mm. Large spec- smooth ventral groove bounds the keel. Rib- imens occasionally have blunt, nodate mid- BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_123 Allen Press • DTPro System File # 01235TQ

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Fig. 104. Prionocyclus quadratus Cobban, 1953. Paratype, USNM 108333c, Sage Breaks Member of the Carlile Shale, sec. 63, T 9S, R 61E, Carter County, Montana. Figures are ϫ1.

¯ank tubercles. Keels are notched with more Ammonites Germari Reuss. SchluÈter, 1872: 41, pl. serrations than ribs. 11, ®gs. 15±17. The bulk of the specimens at hand repre- Ammonites Germari Reuss. Fritsch, 1872: 29, pl. sent the robust form. The gracile form tends 14, ®gs. 1, 2; pl. 16, ®g. 7. to be more densely ribbed and lacks mid¯ank Ammonites SchloÈnbachi Fritsch, 1872: 33, pl. 16, swellings or bullae. Outer ventrolateral tu- ®g. 5. bercles seem to be absent on both robust and SchloÈnbachia Germari Reuss. Fritsch, 1893: 74, ®g. 50. gracile forms. Germariceras germari (Reuss). Breistroffer, 1947: OCCURRENCE: Upper Turonian Scaphites unpaginated. nigricollensis and S. corvensis zones of the Germariceras germari (Reuss). Wright, 1957: northern Great Plains of south-central and L427, ®g. 547.2. southeastern Montana and western South Da- Prionocyclus carvaholi Howarth, 1966: 224, pl. kota, and in equivalent strata in west-central 1, ®gs. 8±11; pl. 2, ®gs. 3±6. New Mexico. Prionocyclus quadratus Cobban. Scott et al., 1986: ®g. 6K. Prionocyclus germari (Reuss, 1845) Prionocyclus germari (Reuss). Kaplan, 1988: 14, Figures 108A, B, D±F; 109±119 pl. 3, ®gs. 1±3, pl. 6, ®g. 1. Ammonites Germari Reuss, 1845: 22, pl. 7, ®g. Prionocyclus ?germari Reuss. Hall et al., 1994: 10. 307, ®g. 4J±L. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_124 Allen Press • DTPro System File # 01235TQ

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Fig. 105. Prionocyclus quadratus Cobban, 1953. Paratype, USNM 108333b, Sage Breaks Member of the Carlile Shale, sec. 63, T 9S, R 61E, Carter County, Montana. Figures are ϫ1.

Germariceras germari (Reuss, 1845). Wright, to nodate umbilical tubercles and nodate to 1996: 186, ®g. 142. 1a±c. clavate inner ventrolateral tubercles; weak Prionocyclus germari (Reuss). Wiese, 1997: ®g. outer ventrolateral clavi are present on some 7.4. juveniles. The gracile form is weakly ribbed. Prionocyclus germari (Reuss, 1845). Kaczsrows- DESCRIPTION: Robust and gracile forms can ki, 2000: 244, ®g. 3a. Prionocyclus germari (Reuss, 1845). Reyment be recognized starting from very small di- and Kennedy, 2001: ®g. 2. ameters. In the juvenile robust form (®g. 113A±P, T, U), the coiling is fairly evolute, TYPE: The lectotype, here designated, is with the umbilical wall notched to accom- the original of Reuss, 1845: 22, plate 7, ®g- modate the inner ventrolateral nodes of the ure 10, from the ``Planermergel von Wer- preceding whorl; the umbilicus represents schowitz.'' The original ®gure is reproduced 36±38% of the diameter (table 12) with a here as ®gure 109K. ¯attened wall and broadly rounded umbilical DIAGNOSIS: Moderately to rather widely shoulder. The intercostal whorl section is spaced, generally rounded, prorsiradiate ribs slightly compressed with the greatest breadth on the outer whorls characterize this species. just outside the umbilical shoulder; the costal Primary and secondary ribs tend to be of section is polygonal with the greatest breadth nearly uniform height. The keel is ®nely at the umbilical bullae. There are 16±24 ribs notched with many more serrations than ribs. per whorl between 25 and 60 mm in most The robust form has ribs that support bullate specimens. The ribs arise at the umbilical BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_125 Allen Press • DTPro System File # 01235TQ

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Fig. 106. Prionocyclus quadratus Cobban, 1953. Paratype, USNM 108333a, Sage Breaks Member of the Carlile Shale, sec. 63, T. 9S, R. 61E, Carter County, Montana. Figures are ϫ1.

seam and bear well-developed umbilical bullae that migrate out to an inner ¯ank position at as little as 20 mm diameter. These give rise to broad, blunt, crowded prorsiradiate ribs that are usually evenly developed, with only occasional irregularities. All bear feebly clavate to conical ¯at-topped inner ventrolateral tubercles that are the bases of long septate spines housed in grooves in the umbilical wall of the succeeding whorl, but rarely preserved (®g. 113X, Y). Low, broad, weakening ribs sweep forward and form an acute chevron with the line of the siphonal keel. Fig. 107. Partial external suture of Prionocy- clus quadratus Cobban, 1953. USNM 498451, Feeble outer ventrolateral clavi are present from the Sage Breaks Member of the Carlile up to 15 mm diameter, but are thereafter lost, Shale in the SW¼ sec. 32, T. 50N, R. 66W, Crook and the ribs are bituberculate. The strong si- County, Wyoming. phonal keel is minutely and evenly crenulat- BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_126 Allen Press • DTPro System File # 01235TQ

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ed. Larger specimens (®gs. 113Q±S, V±Y, quadratus generally has stouter quadrate 114±118) vary in strength and coarseness of whorls, and an incipient lateral tubercle when ribbing, with as few as 22 ribs of equal or adult. unequal strength at 90 mm diameter, at which Ammonites schloÈnbachi Fritsch, 1872 (33, size the umbilical bullae migrate well out to pl. 16, ®g. 5) appears to be based on a poorly the inner ¯ank. This coarse, even or uneven preserved crushed external mold of P. ger- ornament extends to the largest diameters mari. A plaster cast taken from the mold, ap- seen. parently the basis of the original ®gure, is The gracile form (®gs. 109±112) has a shown as ®gure 108A. The ¯ank ribs and whorl breadth to height ratio of 0.66±0.90 long ventrolateral spines compare well with between 25 and 60 mm diameter (table 12). the Western Interior example shown in ®gure The coiling is evolute, with U ϭ 35±40% of 113X, Y. Fritsch ®gured an undoubted ju- diameter; the umbilicus is shallow, with a venile P. germari from the same horizon and low wall. Very crowded, even, primary ribs locality (1872: pl. 16, ®g. 7). arise at the umbilical seam and strengthen Prionocyclus carvaholi Howarth, 1966 (p. across the ¯anks with no or only incipient 224, pl. 1, ®gs. 8±11; pl. 2, ®gs. 3±6; ®g. bullae. Ribs are straight to feebly ¯exuous, 108D±F), from the MocËaÃmedes Desert, An- and prorsiradiate on the ¯anks, where occa- gola, appears to be based on robust variants sional long intercalated ribs yield a total of of the present species, even though Howarth up to 70 ribs per whorl. Each rib bears a tiny (1966: 224) described very small ``upper or conical to feebly clavate inner ventrolateral inner ventrolateral tubercles . . . represented tubercle from which a weakening rib projects merely by a raised portion of the rib obliquely in front of the ventrolateral spine.'' forward to form an acute ventral chevron OCCURRENCE: Fairly abundant at the top of with the minutely and evenly serrated si- the Frontier Formation in central Wyoming phonal ridge. Tiny outer ventrolateral clavi (Cobban, 1990), and in the D-Cross Tongue are present up to a diameter of approximately of the Mancos Shale in south-central New 10 mm, but are thereafter lost. From 60±150 Mexico. The species is also present in the mm this even style of ribbing continues, with basal part of the Austin Chalk in the Rio around 36±40 ribs per whorl, which have Grande area of southwestern Texas, where it feeble but variably developed umbilical bul- was reported as P. reesidei (Freeman, 1961). lae and inner ventrolateral clavi. In all of these areas, P. germari is associated The suture is fairly simple, with a mod- with Mytiloides incertus (Jimbo) and Eubos- erately deep lateral lobe (®g. 119). trychoceras matsumotoi Cobban of late Tu- DISCUSSION: Absence of an outer ventro- ronian, Scaphites corvensis/P. germari Zone lateral tubercle distinguishes this species age. The species also occurs in the upper Tu- from Prionocyclus wyomingensis when ronian of Germany, the Czech Republic, and young, while the absence of an umbilical Angola. plus inner lateral tubercle separates it from P. wyomingensis when adult. The very even Prionocyclus pluricostatus, new species and sparser ribbing of P. germani alone Figure 120 serves to distinguish both juveniles and DERIVATION OF NAME: Plurimus (Latin): adults from P. novimexicanus. Prionocyclus most; costatus (Latin): ribbed.

← Fig. 108. A, B, D±F. Prionocyclus germari (Reuss, 1845). A. plaster cast taken from the natural mold of the holotype of Ammonites SchloÈnbachi Fritsch, 1872: 33, plate 16, ®gure 5, from the upper Turonian P. germari Zone, Priesener Schichten near Werschowitz, Laun, Czech Republic. Narodni Mu- seum, Prague, no. 004823. B. No. C16822 in the same collection, from the same unit near Cernovaly in the Czech Republic. C. The holotype of Ammonites albinus Fritsch, 1872: p. 28, plate 6, ®gure 4, from the Mallnitzer Schichten of Wehlowic, near Melnik, in the Czech Republic. D±F. The holotype of Prionocyclus carvaholi Howarth, 1966, from Ponta Grossa, 3.2 km SSE of Ponta des Salinas, 110 km N. of MocËaÃmedes, Angola. Copy of Howarth, 1966: plate 2, ®gures 3, 4, 6. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_128 Allen Press • DTPro System File # 01235TQ

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TYPES: Holotype is USNM 498491a (®g. tribute them to the upper Turonian ammonite 120D); paratypes USNM 498491b (®g. zone of P. germari, but they may represent 120D), 498492 (®g. 120A), 498493 (®g. a separate, as yet uncharacterized uppermost 120B), and 498494 (®g. 120C), all from Turonian ammonite zone. USGS Mesozoic locality D11898, calcareous OCCURRENCE: As for types. shale 4 m below the base of the Fort Hays Limestone Member of the Mancos Shale, up- Genus Prionocyclites Kennedy, 1988 per Turonian, upper part of Prionocyclus ger- TYPE SPECIES: Prionocyclites mite Kenne- mari Zone, NW ,SE , sec. 22, T. 8 S, R. ¼ ¼ dy, 1988: 89, pl. 10, ®gs. 7±11; text-®g. 30. 87 W, Pitkin County, Colorado. DIAGNOSIS: ``Inner whorls with distant bul- DIAGNOSIS: A densely and evenly ribbed late primary ribs separated by groups of Prionocyclus. There are up to 80 ribs per weaker non-bullate primaries and occasional whorl in juveniles at 20 mm diameter, with shorter intercalated ribs, all with a ventrolat- inner ventrolateral clavi and outer ventrolateral clavus. Keel broadly undulose, not ser- eral clavi, if present, lost at a very early rated. Body chamber smooth but for distant stage. Ribs may be single or arise in pairs or bullate primaries and associated constric- groups at the umbilical shoulder, or interca- tions. Aperture constricted. Suture simple, late low on the ¯anks. Larger specimens with with little-incised elements'' (Kennedy, up to 90 ribs per whorl at 50 mm diameter, 1988: 89). arranged as on nuclei, but bullate ribs bear DISCUSSION: Prionocyclites is a progenetic strong inner ventrolateral clavi that are some- dwarf, presumably derived from Prionocy- times linked with weaker clavi or other ribs. clus, the inner whorls of the two sharing DISCUSSION: The types are crushed in many common features (compare ®g. 121 shale, and there is little to add to the diag- and ®g. 51A±I), although the adult body nosis. Prionocyclus pluricostatus n. sp. bears chamber and simple adult suture of Priono- a super®cial resemblance to P. bosquensis cyclites are quite distinctive. It most closely (®g. 76), also known only from crushed resembles Lymaniceras Matsumoto, 1965, specimens in shales. They differ in that P. here interpreted as another progenetic dwarf bosquensis has ribs of variable strength on collignoniceratid. However, Lymaniceras has nuclei, and persistent inner and outer ventro- a minutely serrated keel and lacks the Prion- lateral tubercles. This last character separates ocyclus-like inner whorls and paucicostate, the species from P. wyomingensis (see ®g. constricted body chamber of Prionocyclites 85A±C, F, G), whereas the eveness of ribbing (modi®ed after Kennedy, 1988: 89). separates it from P. macombi (®g. 67) and P. OCCURRENCE: Prionocyclus hyatti Zone of novimexicanus (®g. 88). P. germari (®gs. north-east Texas only. 109, 113) has even ribbing, but specimens are never as densely ribbed as the present Prionocyclites mite Kennedy, 1988 form. A specimen from New Mexico ®gured Figures 121, 122 by Johnson (1903: 139, pl. 1, ®g. 15) as Prionocyclus appears to be as densely ribbed Prionocyclites mite Kennedy, 1988: 89, pl. 10, as our species, but the New Mexico species ®gs. 7±11, text-®g. 30. is more involute. Prionocyclites mite Kennedy. Emerson et al., The types occur with Mytiloides incertus 1994: 211, unnumbered ®gure on p. 213. at a very high level in the Turonian. We at- TYPE: Holotype by monotypy in USNM

← Fig. 109. Prionocyclus germari (Reuss, 1845). A±C. USNM 498462; D±F. USNM 498463; G, H. USNM 498464; I, J. USNM 443834; all gracile forms from locality 26. K. copy of the original ®gure of the lectotype (Reuss, 1845: plate 7, ®gure 10), from the ``Planermergel von Werschowitz.'' L±N. USNM 498457; O±Q. USNM 498453; R, S. USNM 498458; T, U. USNM 220389; V, W. USNM 498459; X±Z. USNM 498465; A1, B1. USNM 498460, all gracile forms from locality 26. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_130 Allen Press • DTPro System File # 01235TQ

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Fig. 110. Prionocyclus germari (Reuss, 1845). A, B. USNM 498466, gracile form from locality 59. C, D. USNM 498467, gracile form from locality 58. E±G. USNM 498468; H, I, USNM 498469; J, K. USNM 498470, all gracile forms from locality 59. L, M. USNM 498471, gracile form from locality 60. N, O. USNM 498472, gracile form from locality 61. P, Q. USNM 498473, from locality 58. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_131 Allen Press • DTPro System File # 01235TQ

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Fig. 111. Prionocyclus germari (Reuss, 1845). A±C. USNM 498474, gracile form from locality 58. D±F. USNM 498475, gracile form from locality 61. G, H. USNM 356922; I, J. USNM 498476, both gracile forms from locality 59. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_132 Allen Press • DTPro System File # 01235TQ

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Fig. 112. Prionocyclus germari (Reuss, 1845). USNM 498477, a gracile form from locality 60. Figures are ϫ1.

→ Fig. 113. Prionocyclus germari (Reuss, 1845). A±C. USNM 443832; D, E. USNM 498478; F±H, USNM 498452; I, J. USNM 498479; K±M. USNM 498480; N±P. USNM 498481; Q±S. USNM 498454; T, U. USNM 498482; V, W. USNM 498483; X, Y. USNM 443835, all robust forms from locality 26. All ®gures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_133 Allen Press • DTPro System File # 01235TQ

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Fig. 114. Prionocyclus germari (Reuss, 1845). A. USNM 498484, robust form from locality 59. B, C. USNM 498485, robust form from locality 59. D, E. USNM 498486, robust form from locality 58. F, G. USNM 498487, robust form from locality 59. H, I. USNM 356920, robust form from locality 60. All ®gures are ϫ1.

420144 (®gs. 121, 122; Table 13) from the ¯attened, subparallel, with broadly round Arcadia Park Formation, Eagle Ford Group ventrolateral shoulders, venter with strong, at USGS locality 22608 (ex Renfro Collec- blunt, undulose siphonal keel. On phragmo- tion), east of power plant on Mountain Creek cone there are twelve bullate ribs per whorl Lake, Dallas County, Texas; Prionocyclus separated by two slightly weaker nonbullate hyatti zone. ribs, giving a total of thirty-three to thirty- DIAGNOSIS: Small species characterized by four ribs per whorl at the umbilical shoulder. its arched venter and prominent keel. Ribs Ribs are narrow, rounded, prorsiradiate, and are sparse, conspicuously prorsiradiate, and straight on the inner ¯ank, ¯exing forward bear oblique ventrolateral clavi. and concave over the outer ¯ank and ventro- DESCRIPTION: ``Holotype and only known lateral shoulder. A few short ribs intercalate specimen is a complete adult retaining traces on the outer ¯ank and all ribs bear a blunt, of original shell. Coiling moderately evolute, obliquely placed ventrolateral clavus. The with shallow umbilicus, umbilical wall low, ribs project forward from the clavus towards rounded. Whorl section compressed with the mid-line of the venter but decline before whorl breadth to height ratio 0.66. Flanks reaching the keel. The low undulations on BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_135 Allen Press • DTPro System File # 01235TQ

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Fig. 115. Prionocyclus germari (Reuss, 1845). A±C, USNM 498488; D±F. USNM 498455, both robust forms from locality 59. All ®gures are ϫ1.

the keel do not correspond to the ribs; they of the body chamber is near-smooth between are far fewer in number. This ornament ex- three very distant ribs that arise from prom- tends on to the ®rst part of the body chamber inent umbilical bullae. These ribs are narrow, but thereafter rapidly declines. The last part rounded, and efface by the ventrolateral BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_136 Allen Press • DTPro System File # 01235TQ

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Fig. 116. Prionocyclus germari (Reuss, 1845). USNM 498489, a robust form from locality 60. Figures are ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_137 Allen Press • DTPro System File # 01235TQ

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Fig. 117. Prionocyclus germari (Reuss, 1845). USNM 498456, a robust form from locality 26. Figure is ϫ1.

shoulder. They are followed by a shallow not approximated at the end of the phrag- constriction, most obvious at the ventrolat- mocone'' (Kennedy, 1988: 89). eral shoulder. A much stronger constriction DISCUSSION: The only ammonites with marks the adult aperture. Suture very simple which Prionocyclites mite Kennedy, 1988, is with broad, little-incised E/L and narrower L; likely to be confused are species of Lyman- BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_138 Allen Press • DTPro System File # 01235TQ

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Fig. 118. Prionocyclus germari (Reuss, 1845). USNM 498490, a robust form from locality 57. Figure is ϫ1. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_139 Allen Press • DTPro System File # 01235TQ

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Fig. 119. Prionocyclus germari (Reuss, 1845). Partial external suture of USNM 498477 from the upper Turonian P. germari Zone D- Cross Tongue of the Mancos Shale at locality 60.

iceras Matsumoto, 1965. The type species, L. planulatum Matsumoto, 1965 (31, pl. 6, ®gs. 1, 2; pl. 7, ®gs. 1±5; pl. 8, ®gs. 1±8; text-®gs. 10±16), is also a progenetic dwarf, but without the distinctive differentiation of primary ribs seen in Prionocyclites and with a ®nely serrated keel. Body chambers are very different. OCCURRENCE: As for type.

Genus Reesidites Wright and Matsumoto, 1954

TYPE SPECIES: By original designation: Barroisiceras minimum (Yabe ms.) Hayasaka and Fukada, 1951: 325, pl. 1, ®gs. 1±4; pl. 2, ®gs. 1±7. DIAGNOSIS AND DISCUSSION: See Matsu- Fig. 120. Prionocyclus pluricostatus n. sp. A. moto, 1965: 61. Paratype USNM 498492; B. Paratype USNM 498493; C. Paratype USNM 498494; D. Holotype OCCURRENCE: Upper Turonian, Japan, Tu- USNM 498491a (left), and paratype USNM nisia, Armenia, Colombia, and New Mexico 498491b (right), all from locality 34. All ®gures in the U.S. Western Interior. are ϫ1. Reesidites minimus (Hayasaka and Fukada, 1951) 326, pl. 4, ®gs. 1±4, from the Upper Turon- Figures 123, 124 ian of the Ikushumbets, region Hokkaido, Ja- pan. Barroisiceras minimum (Yabe ms.). Hayasaka and Fukada, 1951: 325, pl. 1, ®gs. 1±4; pl. 2, ®gs. DIAGNOSIS: A compressed, involute spe- 1±7. cies that attains a diameter of about 100 mm. Reesidites minimus Hayasaka and Fukada, 1951. The narrow cross section has ¯attened con- Cobban and Kennedy, 1988: 66, ®gs. 1±3 (with vergent and fastigiate venter with sharply de- full synonymy). ®ned ventrolateral shoulder. Ornament of Reesidites minimum (Hayasaka and Fukada, low, broad, somewhat ¯exuous primary and 1951). AmeÂdro, 1990: 270, pl. 25, ®gs. 4, 5 secondary ribs, each supporting a low ven- (with synonymy). trolateral clavus and terminating in a mid- Reesidites minimus (Hayasaka and Fukada, 1951). ventral clavus. Primary ribs begin from low Summesberger and Kennedy, 1996: 119, pl. 13, umbilical bullae. ®gs. 17, 19±23; text-®gs. 15, 16. DESCRIPTION: The smaller specimen, TYPE: Holotype, by original designation, is USNM 414511 (®g. 123A, B) is wholly sep- the original of Hayasaka and Fukada, 1951: tate. Coiling is very involute with an umbi- BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_140 Allen Press • DTPro System File # 01235TQ

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Fig. 121. Prionocyclites mite Kennedy, 1988. A±D. Holotype, USNM 420144, from locality 63. Figure A is ϫ1; Figures B±D are ϫ2.

licus to diameter ratio of 0.12; the umbilical der. The ribs arise in pairs from bullae. There wall is somewhat ¯attened. The whorl sec- are both long and short intercalaries and all tion is very compressed, with a whorl are commonly ¯exed back and convex breadth to height ratio of 0.52, the greatest around mid¯ank. All ribs terminate in ven- breadth at the umbilical bullae; the costal trolateral clavi. The siphonal clavi are very whorl section has ¯attened, convergent sides prominent. On the body chamber the whorl and a fastigiate venter. There are three distant section broadens somewhat and the venter umbilical bullae per half whorl. These give rounds. The umbilical bullae decline and rise to pairs of low, broad, prorsiradiate ribs eventually disappear; most of the ribs are that are ¯exed back on the outer ¯anks. They long, extending to the umbilical shoulder as strengthen across the ¯anks, and additional both ribs and striae, while prominent growth intercalaries arise low on the ¯anks to give lines develop. The ventrolateral and siphonal a total of 13 ribs per half whorl. All ribs bear tubercles decline markedly towards the ap- ventrolateral clavi and strong siphonal clavi erture. on the crest of the fastigiate venter so that The suture line has a markedly asymmet- the shell has a strongly crenulate margin ric, bi®d E/L, the ventral half smaller than when viewed from the side. the dorsal; rectangular L that is deeper than

The larger specimen, USNM 414510 (®g. E; asymmetrically bi®d L/U2; and relatively 123C, D), is crushed, and preserves half a broad and bi®d U2 (®g. 124). whorl of body chamber; it appears to be a DISCUSSION: The larger, crushed specimen nearly complete adult with the umbilical closely resembles the holotype, whereas the seam egressing markedly around the last half smaller matches that ®gured by Obata (1965: whorl. There are ®ve umbilical bullae on the pl. 5, ®g. 5). Reesidites elegans Matsumoto last half whorl of the phragmocone, corre- and Inoma, 1971 (in Matsumoto, 1971: 139, sponding to 18 ribs at the ventrolateral shoul- pl. 23, ®gs. 1±3; text-®gs. 5±7) has much ®n- er, more numerous, delicate, and weaker ribs than R. minimus. R. latus Matsumoto and Obata (1982: 82, pl. 6, ®g. 2) is much more evolute, with U/D ϭ 0.40 at maturity and a broader whorl. R. subtuberculatus (Gerhardt, 1897: 156, pl. 3, ®g. 12) has much weaker ¯ank ornament and tubercles. OCCURRENCE: Scaphites ferronensis/Prion- ocyclus wyomingensis Zone of Valencia County, New Mexico and Scaphites whit®el- Fig. 122. External suture of Prionocyclites di Zone of Socorro County, New Mexico; mite Kennedy, 1988. Holotype, USNM 420144, upper Turonian of Japan, Austria, Tunisia, from locality 63. and Armenia. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_141 Allen Press • DTPro System File # 01235TQ

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Fig. 123. Reesidites minimus (Hayasaka and Fukada, 1951). A, B. USNM 414511, from locality 43. C, D. USNM 414510, from locality 51. All ®gures are ϫ1.

ACKNOWLEDGMENTS (University of South Florida, Tampa) criti- cally reviewed an earlier version of the man- We thank R. E. Burkholder (who took uscript. Financial support of Kennedy by the most of the photographs) and the staffs of the Natural Environment Research Council, Geological Collections in Oxford University Royal Society, and Astor Fund (Oxford) is Museum of Natural History and the Depart- gratefully acknowledged. The U.S. Geologi- ment of Earth Sciences, Oxford, and the Di- cal Survey kindly made the collections avail- vision of Paleontology (Invertebrates) at the able for study. American Museum of Natural History for technical assistance. J. M. Hancock (London/ REFERENCES Shaftesbury), U. Kaplan (GuÈtersloh), R. Ma- pes (Ohio University, Athens), and P. Harries Adkins, W. S. 1928. Handbook of Texas Cretaceous fossils. Texas Univ. Bull. 2838: 385 pp. 1931. Some Upper Cretaceous ammonites in western Texas. Texas Univ. Bull. 3101: 35±211. Adkins, W. S., and F. E. Lozo 1951. Stratigraphy of the Woodbine and Ea- gle Ford, Waco area, Texas. In F. E . Lozo and B. F. Perkins (eds.), The Fig. 124. External suture of Reesidites mini- Woodbine and adjacent strata of the mus (Hayasaka and Fukada, 1951). USNM Waco area of central Texas. Fondren 414511, from locality 43. Sci. Ser. 4: 101±169. BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_142 Allen Press • DTPro System File # 01235TQ

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AmeÂdro, F. Chancellor, G. R., W. J. Kennedy, and J. M. Han- 1990. The ammonites. In F. Robaszynski, M. cock Caron, C. Dupuis, F. AmeÂdro, J.-M. 1994. Turonian ammonite faunas from central Gonzalez Donoso, D. Linares, J. Har- Tunisia. Spec. Pap. Palaeontol. 50: 118 denbol, S. Gartner, F. Calandra, and R. pp. Deloffre (eds.), A tentative integrated Cobban, W. A. stratigraphy in the Turonian of central 1951. Colorado Shale of central and north- Tunisia: formations, zones and sequen- western Montana and equivalent rocks tial stratigraphy in the Kalaat Senan of Black Hills. Bull. Am. Assoc. Pet. area: 259±281, In Bull. Cent. Rech. Ex- Geol. 35: 2170±2198. plor. Prod. Elf-Aquitaine 14: 213±384. 1952. Scaphitoid cephalopods of the Colora- AmeÂdro, F., and G. Badillet do Group. Prof. Pap. U.S. Geol. Surv. 1982. Ammonites du Saumurois. In F. Roba- 239: 42 pp. (1951 imprint). szynski, G. AlcaydeÂ, F. AmeÂdro, G. 1953. A new species of Prionocyclus from Badillet, R. Damotte, J.-C. Foucher, S. the Upper Cretaceous Carlile Shale. J. JardineÂ, O. Legoux, H. Manivit, C. Paleontol. 27: 353±355. Monciardini, and J. Sornay (eds.), Le 1955. Some guide fossils from the Colorado Turonien de la reÂgion-type; Saumurois shale and Telegraph Creek formation, et Touraine, stratigraphie, biozonations, northwestern Montana. In Billings sedimentologie: 130±138. In Bull. Geol. Soc. Guidebook, 6th Annu. Field Cent. Rech. Explor. Prod. Elf-Aquitaine Conf., Sweetgrass arch-Disturbed belt, 6: 119±225. Montana, 1955: 198±207. AmeÂdro, F., and J. M. Hancock 1976. Ammonite record from the Mancos 1985. Les ammonites de l'Autoroute Shale of the Castle Valley±Price± ``L'Aquitaine'', France (Turonien et Woodside area, east-central Utah. Santonien). Cretac. Res. 6: 15±32. Brigham Young Univ. Geol. Stud. 22: AmeÂdro, F., C. ColleteÂ, J. Pietresson de Saint-Au- 117±126. bin, and F. Robaszynski 1983. Molluscan fossil record from the north- 1982. Le Turonien supeÂrieur aÁ Romaniceras (Romaniceras) deverianum de l'Aube eastern part of the Upper Cretaceous (France). Bull. Inf. Geol. Bass. Par. 19: seaway, Western Interior. In W. A. Cob- 29±37. ban and E. A. Merewether, Stratigraphy Barbour, E. H. and paleontology of mid-Cretaceous 1903. Report of the State geologist. Nebraska rocks in Minnesota and contiguous ar- Geol. Surv. 1: 258 pp. eas: Prof. Pap. U.S. Geol. Surv. 1253: Basse, EÂ . 1±25 pp. 1946. Sur deux ammonites nouvelles du Con- 1984a. Mid-Cretaceous ammonite zones, iacien du sud-ouest de Madagascar; Western Interior, United States. Bull. Subbarroisiceras n. g. mahafalense n. Geol. Soc. Denmark 33: 71±89. sp. et Eboroceras n. g. magnumbilica- 1984b. Molluscan record from a mid-Creta- tum n. sp. Bull. Soc. GeÂol. Fr. (5) 16: ceous borehole in Weston County, Wy- 71±76. oming. Prof. Pap. U.S. Geol. Surv. Bengston, P. (Compiler) 1271: 24 pp. 1996. The Turonian stage and substage 1986. Upper Cretaceous molluscan record boundaries. Bull. R. Sci. Belg. Sci. Ter- from Lincoln County, New Mexico. re 66(Suppl.): 69±79. Southwest Section of AAPG, transac- Boyle, C. tions and guidebook of 1986 Conven- 1893. A catalogue and bibliography of North tion, Ruidoso, New Mexico: 77±89. American Mesozoic Invertebrata. Bull. 1990. Ammonites and some characteristic bi- U.S. Geol. Surv. 102: 315 pp. valves from the Upper Cretaceous Breistroffer, M. Frontier Formation, Natrona County, 1947. Notes de nomenclature paleÂozoolo- Wyoming. Bull. U.S. Geol. Surv. 1917- gique. P.-V. Mens. Soc. Sci. DauphineÂ, B: 13 pp. 26th year, no. 195: 5 pp. (unnumbered). Cobban, W. A., and S. C. Hook Case, G. R. 1979. Collignoniceras woollgari woollgari 1982. A pictorial guide to fossils. New York: (Mantell) ammonite fauna from Upper Van Nostrand Reinhold, 515 pp. Cretaceous of Western Interior, United BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY Tuesday Nov 20 2001 04:36 PM 2000 amnb 01235 Mp_143 Allen Press • DTPro System File # 01235TQ

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