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Novtautesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y NovtautesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2722, pp. 1-24, figs. 1-1I1 January 29, 1982 Studies on the Paleozoic Selachian Genus Ctenacanthus Agassiz: No. 2. Bythiacanthus St. John and Worthen, Amelacanthus, New Genus, Eunemacanthus St. John and Worthen, Sphenacanthus Agassiz, and Wodnika Miunster JOHN G. MAISEY1 ABSTRACT Some of the finspines originally referred to Eunemacanthus St. John and Worthen is revised Ctenacanthus are reassigned to other taxa. Sev- to include some European and North American eral characteristically tuberculate lower Carbon- species. Sphenacanthus Agassiz is shown to be iferous finspines are referred to Bythiacanthus St. a distinct taxon from Ctenacanthus Agassiz, on John and Worthen, including one of Agassiz's the basis of finspine morphology, and its wide- original species, Ctenacanthus brevis. Finspines spread occurrence in the Carboniferous of North referable to Bythiacanthus are known from west- America is demonstrated. Similarities are noted ern Europe, the U.S.S.R., and North America. between the finspines of Sphenacanthus and Amelacanthus, new genus, is described on the Wodnika, and both taxa are placed provisionally basis of finspines from the United Kingdom. Four in the family Sphenacanthidae. A new species of species are recognized, two of which were origi- Wodnika, W. borealis, is recognized on the basis nally assigned to Onchus by Agassiz, and all four of a finspine from the Permian of Alaska. of which were referred to Ctenacanthus by Davis. INTRODUCTION The present paper is the second in a series Ctenacanthus in an attempt to restrict this of reviews of the Paleozoic chondrichthyan taxon to sharks with finspines that closely Ctenacanthus. The first paper (Maisey, resemble those of the type species, C. ma- 1981) reexamined Agassiz's (1837) genus jor. Agassiz (1837) described some other 1 Assistant Curator, Department of Vertebrate Paleontology, American Museum of Natural History. Copyright (©) American Museum of Natural History 1982 ISSN 0003-0082 / Price $2.00 2 AMERICAN MUSEUM NOVITATES NO. 2722 spines which were referred to Ctenacanthus, different from that of C. major in shape and although their morphology and ornament ornamentation. Instead of being long and patterns differ greatly from those of C. ma- slender, the spine of C. brevis is squat and jor. Only one of Agassiz's (1837) other Cten- broad, and is ornamented by large round, acanthus species, C. tenuistriatus, is left in striated tubercles arranged in closely spaced that genus by Maisey (1981). However, var- series down the spine, instead of the fine pec- ious spines have been referred to Ctenacan- tinated ribs seen on C. major spines. Since thus by previous authors, with the result that the spines of C. brevis and C. major are so the genus had become reduced to an almost different, retaining C. brevis in the genus undiagnosable state; in particular, Sphen- Ctenacanthus is unjustified. Recent squaloid acanthus has been greatly confused with and heterodontid finspine morphology and Ctenacanthus. The present work therefore ornamentation does not vary below generic reviews finspines that were included in Cten- level. This is apparently also true of hybo- acanthus by Agassiz (1837) but were exclud- dont finspines (Maisey, 1978). ed by Maisey (1981), and also includes a re- Agassiz (1837) never saw the finspine on view of the genus Sphenacanthus. which his description of C. brevis was based before its publication, although it is known ACKNOWLEDGMENTS which specimen this was and it can still be I thank the following people for their as- located (see below and fig. 2B, C). He knew sistance in preparing this work: Ms. Lorraine the specimen only from a drawing sent by Meeker and Mr. Chester Tarka (preparation the Reverend William Buckland, his col- of illustrations), Mr. Walter Sorensen (clean- league. ing and preparation of specimens). The fol- Finspines similar to those in C. brevis lowing have also been most helpful in were later described from North America providing specimens, photographs or and referred to a new genus, Bythiacanthus information; Drs. S. M. Andrews, M. D. St. John and Worthen (1875). Comparison of Crane, C. L. Forbes, W. J. Hlavin, N. Hot- the original descriptions with other finspines ton, D. Sartenaer, B. Schaeffer, G. Schaum- referred to Bythiacanthus in the American berg, K. S. Thomson, M. Williams, R. Zan- Museum of Natural History suggest that gerl and J. Zidek. Abbreviations are: AMNH, Agassiz's (1837) C. brevis should provision- American Museum of Natural History; ally be referred to Bythiacanthus. BMNH, British Museum (Natural History); AMENDED DIAGNOSIS: Elasmobranch rec- CM, Carnegie Museum; USNM, United ognized by having dorsal finspines of stout States National Museum, Smithsonian In- build, rhomboidal outline in lateral view; stitution. thick walls of trabecular osteodentine; ante- rior face rounded, strongly to moderately compressed laterally, posterior wall convex, BYTHIACANTHUS ST. JOHN AND WORTHEN level of posterior closure very high and the HISTORICAL NOTE: The type species of posterior wall correspondingly short; in Ctenacanthus (Agassiz, 1837) is C. major. transverse section the trunk wall much thick- The second species he described, C. brevis, ened anteriorly; ornament of longitudinal is based on a dorsal finspine which is very rows of rounded, striated tubercles, usually FIG. 1. A-I, Bythiacanthus vanhornei; holotype, from St. John and Worthen (1875) pl. 17, no. 1. J-L, Bythiacanthus siderius (Leidy); J-K, AMNH 1826, St. Louis Limestone, Alton, Illinois; L, the holotype PAN S 22:17 7835, "Sub-Carboniferous," Glasgow, Tennessee; from Leidy (1873) pl. 32, no. 59. M-N, Bythiacanthus off. ianishevskyi AMNH 9594 (reversed to facilitate comparison with other specimens); Waverly sandstone, ?Marion Co., Kentucky. 1982 MAISEY: CTENACANTHUS AGASSIZ 3 A B ~~~~~~~~~~~~~~~~~~~~~~~J\ 2mm 2cm G H M N 4 AMERICAN MUSEUM NOVITATES NO. 2722 B 1 C\j~~~~~~~~~~~~~~~~~~~~~~- 2mm G J~~~~~~L I- i'e s / E K 2mm L FIG. 2. A-F, Bythiacanthus brevis (Agassiz); A, from Agassiz (1837) tab. 2, no. 2; B, C, the holotype C 4154 (photographs courtesy of Bristol City Museum and Art Gallery, England). Compare with A which was originally copied from Buckland's drawing of this specimen; D, deta-il of ornament from another specimen, BM(NH) P2226; E, F, from Davis (1883) p1. XLIII, no. 3; this appears to be specimen BM(NH) P2226. G-K, Bythiacanthus ianishevskyi (Khabakob); from Khabakob (1926) pl. III, nos. 5-10. 1982 MAISEY: CTENACANTHUS AGASSIZ 5 C Ev- F a F-r D E ., -.: i FIG. 3. A, B, Bythiacanthus solidus (Eastman); A, the holotype, USNM 3383; B, paratype, from Eastman (1903) p1. 7, no. 3. C-F, Bythiacanthus peregrinus (Khabakob); from Khabakob (1928) pl. III, nos. 1-4. G, Bythiacanthus lucasi (Eastman); from Eastman (1902) pl. 6, no. 1. less than their own diameter apart; rows in- Asteracanthus siderius Leidy, 1873, p. 313; creased proximally by primary bifurcation C. solidus Eastman, 1902, p. 90; Incertae anteriorly and by being inserted between sedis, aff. Bythiacanthus; Glymmatacan- other rows marginally. thus irishii St. John and Worthen, 1875, p. TYPE SPECIES: Bythiacanthus vanhornei 447; G. rudis St. John and Worthen, 1883, St. John and Worthen, 1875, p. 445, St. p. 249; G. petrodoides St. John and Wor- Louis Limestone, Alton, Illinois. then, 1883, p. 250. REFERRED SPECIES: Ctenacanthus brevis DISCUSSION: Leidy (1873, p. 313) referred Agassiz, 1837, p. II; C. ianishevskyi Kha- a coarsely tuberculated finspine, PAN S bakob, 1928, p. 23; C. lucasi Eastman, 1902, 22:13 7835, from the "Sub-Carboniferous" p. 80; C. peregrinus Khabakob, 1928, p. 25; of Glasgow, Tennessee, to a new species, 6 AMERICAN MUSEUM NOVITATES NO. 2722 Asteracanthus siderius. Apart from the pres- relatively even deeper cross-sections are ence of tubercles, however, there is nothing also known. to suggest that this spine is referable to As- Bythiacanthus solidus (Eastman, 1902, p. teracanthus, which is otherwise a Mesozoic 90) is based on finspines USNM 3383 (the taxon and which has distinctive hybodontid holotype) and USNM 4833 (paratype). East- finspines (Maisey, 1978). Leidy's (1873) man's (1902, fig. 13) transverse section is specimen of a spine resembles that of By- taken near the base of the type finspine, but thiacanthus vanhornei, described by St. does not indicate the full depth of the spine John and Worthen (1875, p. 445), who sug- anteroposteriorly. Eastman (1902, pl. 2, fig. gested that these spines might belong to the 3) did not otherwise figure the type speci- same species. Another almost complete fin- men, which is illustrated here (fig. 3A). The spine, AMNH 1826, closely resembles Lei- ornament of the holotype is slightly less pro- dy's (1873) specimen, and is referred to By- nounced than that of the paratype (fig. 3B), thiacanthus siderius. Finspines referred to and is similar to the ornament of B. lucasi, this taxon are more densely covered by tu- also described by Eastman (1902, p. 80, pi. bercles than those in B. vanhornei. Because 6, fig. 1, also text-fig. 9). The ornamentation of the differences in their tubercle arrange- of B. solidus, however, is less regular than ment and density, I disagree with St. John that of B. lucasi (fig. 3G). A primary row of and Worthen's (1875) suggestion that these tubercles is present anteriorly on finspines of species are synonymous (fig. lA-L). both species. Both are provisionally referred Agassiz's (1837) figure of "Ctenacanthus" to Bythiacanthus although their ornamenta- brevis is reproduced here (fig. 2A), along tion is more regular than in the type species, with illustrations of the actual specimen (fig. B. vanhornei. 2B, C). This represents the first time that the Bythiacanthus ianishevskyi (Khabakob, type specimen has been properly figured. 1928, p. 23) is also founded on a fragmentary The coarsely tuberculate ornament is shown finspine with coarse, tuberculate ornament in detail (fig.
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