Novltates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

AMERICANt MUSEUM Novltates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2774, pp. 1-20 January 27, 1984

Studies on the Paleozoic Selachian Genus Ctenacanthus Agassiz. No. 3. Nominal Species Referred to Ctenacanthus

JOHN G. MAISEY'

ABSTRACT All known nominal species of the Paleozoic another spine genus, Acondylacanthus; three are chondrichthyan genus Ctenacanthus are listed, to- referred to Asteroptychius; nine are referred to gether with information concerning provenance, Sphenacanthus; 20 are referred to various other whereabouts of type material (where known), im- genera; eight are considered close to Ctenacanthus portant subsequent references and comments on but are excluded from it; and 23 are left in Cten- the probable affinities of each species. Other ref- acanthus. Of the latter, however, eight are prob- erences to undetermined Ctenacanthus spp. are ably synonyms of the type species, C. major, and listed chronologically. Ofsome 100 species, 10 are consequently only 15 species are retained. Ofthese, considered totally invalid; the holotypes of five seven are from the Upper Devonian and eight are others are known to be lost or destroyed; four from the Lower Carboniferous; all are from marine others are founded on inadequate material; 10 are strata. This stratigraphic range is much less than referred to the Acanthodii; eight are referred to the previous records have suggested.

INTRODUCTION In the two preceding parts of this series future. In the meantime, it seems advisable (Maisey, 1981, 1982), the diagnosis of Cten- to publish a compilation of data concerning acanthus has been refined to agree with the prior references to Ctenacanthus finspines. type species, C. major, and a list of species The checklist and references given below rep- founded on finspines similar to those of C. resent the majority ofpublished accounts. Al- major has been published. Certain other fin- though I have tried to make this list as com- spines, differing strongly from those of C. plete as possible, perhaps it is inevitable that major, were referred to various genera in part a few sources have escaped notice. Never- two of these studies. Many additional forms theless, no similar listing has been compiled remain to be surveyed, and it is hoped that since the publication of the second volume some of these will be reviewed in the near of Woodward's (1891) Catalogue, and the

' Assistant Curator, Department of Vertebrate Paleontology, American Museum of Natural History.

Copyright © American Museum of Natural History 1984 ISSN 0003-0082 / Price $1.70 2 AMERICAN MUSEUM NOVITATES NO. 2774 present compilation may therefore be useful Ctenacanthus abnormis Giebel (1858, p. 264, to paleontologists, zoologists, and biostratig- pl. 1, fig. 12). raphers. Where the location of a type speci- Lower Devonian, Harz Mountains. men is known, it is given, together with its COMMENTS: See also Barrande (1872, p. catalogue number. Where this datum is ab- 628), Kayser (1878, p. 3, Taf. 1, fig. 19). Re- sent, the whereabouts of the type material is ferred to Machaeracanthus by Woodward unknown (although this does not necessarily (1891, p. 123). mean that it is lost, only that I have not been able to locate it). In order to increase the Ctenacanthus acutus Eastman (1897), p. 12, usefulness ofthe data, notes have been added fig. 2). after each species is listed, and the affinities Type specimen, USNM 4683; Mississip- and validity of the species are discussed. pian, Keokuk Limestone, Iowa. Species of Ctenacanthus are listed in alpha- COMMENTS: Another specimen, USNM betical order, and are followed by a list of 4682 may be referred to this species. In trans- indeterminate references to "Ctenacanthus verse section and ornamentation pattern C. sp." acutus finspines resemble those of Tristy- I stress that, while many species have been chius. USNM 3480 said to be close to C. referred to Ctenacanthus, only a small num- acutus (Eastman, 1902, p. 83). ber are based on finspines that compare Ctenacanthus aequistriatus Davis (1879a, p. closely with the type species. Many of the 185, pl. X, fig. 5). species herein have subsequently been re- Lower Coal Measures (Pennsylvanian), ferred to other genera, and probably even Yorkshire, England. more of them will be removed as work pro- COMMENTS: This form was subsequently gresses. The reader is cautioned here, and is referred to the genus Sphenacanthus by reminded in text, that some of these spines Woodward (1889, p. 244) and Maisey (1982). may pertain to very different fishes from Ctenacanthus amblyxiphias Cope (1891, p. Ctenacanthus. For diagnoses and discussions 449, pl. XXVIII, fig. 3). of the genus, the following are the main Type 7289, Permian, sources: Agassiz (1837), McCoy (1855), New- specimen, AMNH Davis Texas. berry (1873), De Koninck (1878), COMMENTS: Other references include East- (1883), Woodward (1891), Eastman (1902, man (1903), Woodruff (1906), Hussakof 1907), Obruchev (1967), Maisey (1981, (1908, 1911), Branson (1916), Romer(1942), 1982). Wells (1944), Berman (1970), and Zidek INSTITUTIONAL ABBREVIATIONS (1976). According to Wells (1944), however, AMNH, American Museum of Natural History Eastman's (1903) finspine is closer to that of BM(NH), British Museum (Natural History) C. lamborni; this is also the case with Bran- CNIGR, Central Scientific Research, Geological son's (1916) specimen. Prospecting Museum, Leningrad MCZ, Museum ofComparative Zoology, Harvard Ctenacanthus angulatus Newberry and Wor- University then (1866, p. 118, pl. XII, fig. 4). USNM, National Museum of Natural History, Type specimen, University of Illinois Smithsonian Institution, Washington, D.C. X- 1152, Mississippian, Chester Limestone, YPM, Yale Peabody Museum Chester, Illinois. ALPHABETICAL LIST OF SPECIES COMMENTS: There are similarities between C. angulatus and C. similis finspines, but All the following species have been referred these are not sufficient to suggest synonymy. to Ctenacanthus, in most cases by the original Ctenacanthus angulatus is among the species author, but occasionally by a subsequent one. retained in the genus by Maisey (1981). The name is given, followed by the author, reference date, page and illustration num- Ctenacanthus angustus Newberry (1889, p. bers. If known, the type specimen is listed, 181). with locality data if any. Additional data are Type specimen, AMNH 5269, "Subcar- then given under Comments. boniferous," Berea Grit, Berea, Ohio. 1984 MAISEY: CTENACANTHUS AGASSIZ 3

COMMENTS: The type was not figured by Ctenacanthus and referred instead to Bythi- Newberry (1889) but the specimen so des- acanthus by Maisey (1981, 1982). Several ignated bears a label in his handwriting. A other specimens were listed by Woodward detail of its ornament was subsequently fig- (1891, p. 100), who noted that C. limaformis ured (Hussakof, 1908, fig. 19). The species is is probably synonymous with this species. among those retained in the genus by Maisey (1981). Some specimens referred to "C. an- Ctenacanthus browni Branson (1916, p. 653, gustens" e.g., BM(NH) P9581, P9262, may pl. IV, fig. 7, text fig. 6). pertain to this species. Type specimen, University ofMissouri no. 709, Upper Carboniferous, Embar Forma- Ctenacanthus bellus Branson (1906, p. 1393, tion, Wyoming. pl. XLII, figs. 19-21). COMMENTS: An additional specimen was Syntypes, AMNH 6446 (two pieces), Mis- referred to the species by Branson (1933). sissippian, Salem Limestone, Lanesville, In- The finspine was not included in Ctenacan- diana. thus by Maisey (1981); it is more probably COMMENTS: This species is not referable to allied to Acondylacanthus. Ctenacanthus. The wedge-shaped transverse section, tubercles toward the leading edge, Ctenacanthus burlingtonensis St. John and and presence of a narrow median ridge pos- Worthen (1875, p. 426, pl. XV, figs. 6, 7). teriorly characterize this as an Asteroptychius Cotypes, USNM 13523 and 13524, Mis- finspine. sissippian, St. Louis Limestone, St. Louis, Missouri. Ctenacanthus bohemicus Barrande (1872). COMMENTS: An additional specimen, Lower Devonian (Siegenian), Lochkov AMNH 1031, seems close to the type ma- Limestone (Bohemia), Taunusquartzit terial in many respects. The fragments upon (Rheinland). which this species is based are remarkably COMMENTS: Referred to Machaeracanthus similar to finspines of Acondylacanthus gra- by Woodward (1891, p. 124), Fritsch (1895, cillimus. Probably C. burlingtonensis is syn- p. 72), and Denison (1979, p. 52); probably onymous with C. gradocostus. an acanthodian. Ctenacanthus buttersi St. John and Worthen Ctenacanthus bosnensis Katzer (1916, p. 201, (1883, p. 240, pl. XXII, fig. 2). Tab. 1, figs. 1-13), Pennsylvanian, Lower Coal Measure, Car- Triassic, near Sarajevo, Bosna i Hercego- linville, Illinois. vina, Yugoslavia. COMMENTS: The ornamentation ofthe type COMMENTS: This species is probably refer- specimen differs from that of Ctenacanthus able to Nemacanthus. Its finspines are char- finspines. The general shape and the arrange- acteristically ornamented with longitudinal ment ofthe ornament in C. buttersi are rem- series of rounded tubercles laterally and an iniscent ofMesozoic hybodont finspines. This enameled keel anteriorly. form does not seem close to Sphenacanthus. A spine from the Seminole Formation of Ctenacanthus brevis Agassiz (1837, vol. III, Oklahoma was said to be reminiscent of C. p. 11, table 2, fig. 2). buttersi by Zidek (1977, p. 153). Type specimen, Bristol City Museum "Ctenacanthus buttlersi St. John and Wor- C4154, Mississippian, Carboniferous Lime- then"; (in) Caster (1930, p. 100), a typograph- stone (Avonian, Z2 fishbeds), Clifton, near ical error for C. buttersi (see above). Bristol, England. COMMENTS: Agassiz (1837) figured a draw- Ctenacanthus cannaliratus St. John and ing rather than the specimen, which he had Worthen (1883, p. 239, pl. XXI, fig. 3). not seen prior to publication. The type spec- Type specimen, University of Illinois imen is figured in Maisey (1982, fig. 2). X-391, Mississippian, Chester Limestone, Another specimen, BM(NH) 3111-2, was de- Chester, Illinois. scribed and figured by Davis (1883, p. 337, COMMENTS: The fragment upon which this pl. XLIII, fig. 3). The species is removed from species is based differs from Acondylacanthus 4 AMERICAN MUSEUM NOVITATES NO. 2774 gracillimus only in the greater delicacy of its Maisey (1981), who also included Dean's costae and tubercles, and C. cannaliratus is (1909) specimen of"C. clarkii" in the present considered synonymous with A. gracillimus species, which is retained in the genus. (see also C. gracillimus, below). Ctenacanthus ? costatus Newberry and Wor- Ctenacanthus chemungensis Eastman (1907, then (1866, p. 120, pl. XII, fig. 5). p. 77, pl. 7, fig. 3). Mississippian, St. Louis Limestone, St. Devonian, Chemung Group, Bradford Co., Louis, Missouri. Pennsylvania. COMMENTS: The broad, smooth and heavi- COMMENTS: The species was originally ly enameled costae of the type specimen are named by Claypole (1885, p. 490). Eastman atypical of Ctenacanthus finspines. Subse- (1907) figured a detail of ornament from an quently this form was made the type species unspecified specimen (reproduced by Caster, of Eunemacanthus (St. John and Worthen, 1930, pl. 56, fig. 9), and no type specimen 1883, p. 246, pl. XXIII, fig. 2; see also Mai- was designated; from the literature it ought sey, 1982). to be one of the specimens collected by Ctenacanthus costellatus Traquair (1884, p. Beecher and deposited at New Haven or 3, pl. II). Cambridge. Eastman (1907) referred to ad- Type specimen, BM(NH) P5900, Missis- ditional specimens collected in New York; sippian, Calciferous Sandstone, Dumfries- see also Caster (1930, p. 102). shire, Scotland. Ctenacanthus clarkii Newberry (1889, p. 168, COMMENTS: This species is known from a pl. XXVI, figs. 2, 3). complete, articulated specimen, subsequent- Type specimen, AMNH 220G, Cleveland ly described by Moy-Thomas (1936, p. 761). Shale, Berea, Ohio. Its finspines do not agree with those of Cten- COMMENTS: See also Hussakof (1908). A acanthus, however, and the species has been partial skeleton, with teeth, jaws, pectoral fins, referred provisionally to Sphenacanthus shagreen and part of the impression of a fin- (Woodward, 1889; Maisey, 1982). spine was referred to C. clarkii by Dean (1909). According to Maisey (1981) Dean's Ctenacanthus coxianus St. John and Wor- specimen should more correctly be referred then (1883, p. 233, pl. XXI, fig. 1). to C. compressus, but C. clarkii and C. com- Type specimen, USNM 13502, Mississip- pressus are retained in the genus as separate pian, Keokuk Limestone, Montrose, Iowa; species. plesiotype, MCZ 5188. COMMENTS: Eastman (1902, p. 87) noted Ctenacanthus cliftonensis Branson and Mehl considerable resemblance between finspines (1938, p. 122, pl. 37, figs. 16-19). ofthis species and those of C. furcicarinatus, Type specimen, University of Missouri, and they may be closely related. An almost 752 VP, lower Mississippian, Chouteau complete finspine, AMNH 11201 (Linney Limestone, Clifton City, Missouri. Coll.), lacking only the apex was collected COMMENTS: This form is similar to C. va- supposedly from the Waverly Series, Marion rians, but is provisionally retained as a sep- Co., Kentucky. arate species of Ctenacanthus (Maisey, 1981). Ctenacanthus crenatus McCoy (1855, p. 624, Ctenacanthus compressus Newberry (1889, pl. 3.1., fig. 31). p. 168, pl. XXII, fig. 4). Mississippian, Carboniferous Limestone, Type specimen, AMNH 140G, Cleveland Armagh. Shale, Sheffield, Lorain Co., Ohio. COMMENTS: According to McCoy (1855) COMMENTS: The species was first named and Davis (1883), this species is the same as and briefly described some years earlier that named C. crenulatus by Agassiz (1837, (Newberry, 1878, p. 191), but it was not fig- vol. III, p. 177, name only). McCoy's (1855) ured in that work. The figure appearing in figure was reproduced by Davis (1883, pl. 1889 is reversed. See also Hussakof (1908). XLV, fig. 6), but was reversed. Davis noted The type and other specimens are figured by that the type specimen (a fragment of spine) 1 984 MAISEY: CTENACANTHUS AGASSIZ 5 was lost; it has never been relocated. Both Ctenacanthus denticulatus McCoy (1855, p. species, C. crenatus and C. crenulatus should 625, pl. 3K, fig. 16). therefore be declared nomina nuda. Mississippian, Carboniferous Limestone, Ctenacanthus crenulatus Agassiz (1837, vol. Armagh and Drumlish (Northern Ireland), III, p. 177; name only). Shropshire, England. See C. crenatus (above). COMMENTS: The species was named but not figured by McCoy (1848, p. 116). See also Ctenacanthus cylindricus Newberry (1889, p. Davis (1883, p. 338, pl. XLIV, fig. 4) and 202, pI. XXVI, fig. 1). Maisey (1981, fig. 8F, G). It is retained in Type specimen, AMNH 358G, "Subcar- Ctenacanthus by Maisey (1981). boniferous," Keokuk Group, Casey Co., Kentucky. Ctenacanthus depressus Newberry (1897, p. COMMENTS: It is doubtful whether this 291, pl. XXII, fig. 6). species is based on a median (unpaired) spine. Mississippian, Kinderhook Group, Le In transverse section and style or ornamen- Grand, Iowa. tation the spine resembles those of Gyracan- COMMENTS: This form may be close to C. thus, except that the costae are less strongly buttersi, as Newberry (1897) suggested. Its arranged into chevrons. I do not regard this smooth ornamentation pattern seems to pre- as a Ctenacanthus species. clude it from the genus Ctenacanthus, and it may be closer to Sphenacanthus. Ctenacanthus decussatus Eastman (1902, p. 84, pl. 6, fig. 2, text-fig. 11). Ctenacanthus distans McCoy (1855, p. 625, Type specimen, USNM 4846, Mississip- pl. 3K, fig. 15). pian, Kinderhook Limestone, location un- Mississippian, "red limestone" ofArmagh, certain (Iowa or Illinois). Northern Ireland. COMMENTS: The type specimen resembles COMMENTS: These elongate, slender spines that of C. buttersi, but not C. pellensis (cf. resemble those of C. formosus, but more par- Eastman, 1902). Its ornamentation pattern, ticularly those of Acondylacanthus gracilli- in my view, precludes it from the genus Cten- mus. Originally named but not figured by acanthus. McCoy (1848, p. 116), the species was subsequently transferred to Acondylacanthus by Ctenacanthus deflexus St. John and Worthen Davis (1883, p. 349) and Woodward (1891, (1883, p. 234, pl. XXII, fig. 1). p. 108). Mississippian, St. Louis Limestone, Alton, Illinois. Ctenacanthus dubius Davis (1883, p. 340, pl. COMMENTS: St. John and Worthen (1883) XLIV, fig. 7). regarded this species as being allied to C. Type specimen, BM(NH) P2530, Missis- speciosus, which is itself probably synony- sippian, "Mountain Limestone" of Armagh, mous with C. varians (Eastman, 1902; Mai- Northern Ireland. sey, 1981). Maisey (1981) retained them as COMMENTS: This species is founded on a separate species within the genus Ctenacan- badly crushed and distorted finspine. Its or- thus, but suggested that C. deflexus may be namentation is atypical of Ctenacanthus and synonymous with C. major. the species is provisionally referred to Ame- lacanthus (see Maisey, 1982). Ctenacanthus deliculatus (Eastman), Deni- son (1979, p. 52). Ctenacanthus elegans Tuomey (1858, p. 38, COMMENTS: The species was removed from woodcut, fig. A). Homacanthus by Denison, who suggested that Devonian; base of Chattanooga Shale, it may instead pertain to Ctenacanthus. See Shoal Creek, nr. Florence, Lauderdale Co., Eastman (1903, p. 218, pl. III, fig. 28, and Alabama. pl. V, fig. 59). The holotype of H. deliculatus COMMENTS: Apparently referable to Cten- is MCZ 5126. It is not referable to Cten- acanthus (Tuomey suggested that "it may turn acanthus however, and its affinities are un- out to be identical with C. tenuistriatus"). certain. Unfortunately, the type specimen is now lost. 6 AMERICAN MUSEUM NOVITATES NO. 2774

I am indebted to Dr. Charles Copeland (Geo- its long, elegant slender finspines. See also logical Survey of Alabama) for information Newberry (1875, p. 53, pl. LIX, fig. 1). Not concerning this specimen. It was probably of listed as a member of Ctenacanthus by Mai- late Devonian rather than Mississippian age sey (1981) but probably close to this genus. as thought by Tuomey. Apparently all of The holotype was lost for many years, but Tuomey's records and specimens were lost through the endeavors ofRobert Hook (Uni- sometime between his death (1857) and the versity of Kentucky) it has recently been re- appointment in 1873 ofhis successor. During located (March 1983) along with a significant this interim the University ofAlabama cam- part ofthe original William Patterson collec- pus was destroyed by the 2nd Michigan Cav- tion (formerly University of Kentucky UK alry (April 4 and 5, 1865), and the type spec- 1004 = M2060). The holotype has now been imen is unlikely to have survived! transferred to the American Museum ofNat- designation above). Ctenacanthus? erectus v. Koenen (1895, p. ural History (see 3). Ctenacanthusfurcicarinatus Newberry (1875, Upper Devonian, Biidesheim bei Gerol- p. 54, pl. LIX, fig. 2). stein. Type specimen, AMNH 11543, Famen- COMMENTS: According to Gross (1933, p. nian, Waverly Black Shale, Vanceburg, Ken- 65) this finspine has ornament like that of tucky. Onchus curvatus from the upper Silurian. The COMMENTS: The finspine ornamentation species may therefore represent an acantho- pattern is atypical for Ctenacanthus. The dian, and is not regarded here as a member species is of considerable interest, since its of Ctenacanthus. The specimen has not been finspines were found in association with teeth figured. of"Orodus variabilis" and pieces of shagreen with a regular, gridlike squamation pattern Ctenacanthus excavatus St. John and Wor- of ridged scales, e.g., AMNH 409. Newberry then (1875, p. 428, pl. XV, figs. 4, 5). (1875) also mentioned that fairly complete Mississippian, Iowa and Missouri. sharks had once been found by quarrymen COMMENTS: The species is founded upon but none of these specimens has ever been the apical fragments of two finspines (possi- described. The holotypes of C. furcicarinatus bly juvenile). It is difficult to assign these and 0. variabilis, together with a tail of a pieces to Ctenacanthus; they could represent shark mentioned by Newberry (1875) as pos- small Eunemacanthus finspines (see C. cos- sibly pertaining to C. furcicarinatus have been tatus). rediscovered (1983) by Robert Hook and Ctenacanthus? fallax Leidy (1857, pl. V, fig. transferred to the American Museum. 30). Ctenacanthus gemmatus St. John and Wor- Type specimen, Museum of the Academy then (1875, p. 429, pl. XV, figs. 9, 10). of Sciences, Philadelphia, no. 7798. Missis- Mississippian, St. Louis Limestone, Alton, sippian, St. Louis Limestone, St. Louis, Mis- Illinois. souri. COMMENTS: Species is probably referable COMMENTS: "founded on an indetermina- to Acondylacanthus; it is fairly similar to A. ble fossil" (Woodward, 1891, p. 105). How occidentalis. true! Illustrated but not named or described by Leidy (1857). See Gillette and Shapiro Ctenacanthus gemundensis Gross (1933, p. (1978, p. 116). 64, p1. II, fig. 8). Type specimen, Geologische Landesan- Ctenacanthus formosus Newberry (1873, p. stalt, Berlin. 328, pl. 36, fig. 2). Late Devonian, Schliedener Schichten, Holotype, AMNH 11544, Famennian, Gemuind (Eifel). Waverly Series, Vanceburg, Kentucky; also COMMENTS: An indeterminate finspine Berea Grit and Cuyahoga Shale (e.g., AMNH founded on very fragmentary material. Or- 221, 223) of Ohio. nament pattern atypical ofCtenacanthus; may COMMENTS: A species easily recognized by be an acanthodian finspine; referred to No- 1 984 MAISEY: CTENACANTHUS AGASSIZ 7 dacosta, Gross (1940) by Denison (1975, p. species is founded on indeterminate material. 53). The type specimen was apparently de- Elsewhere in the present work I have made stroyed in World War II (Dr. K. Fischer, per- certain species synonymous with A. gracil- sonal commun.). limus rather than with A. occidentalis, which to all intents can be left as a separate, inde- Ctenacanthus gondwanus Silva Santos (1947, terminate species. See also C. cannaliratus. p. 248, pl. 1, figs. 1-5). The Ctenacanthus sp. finspine of Kul- Type specimen, National Geological Mu- czycki (1957, pl. XIII, fig. 2) may be referable seum, Brazil, DGM 73-P. Pennsylvanian, to A. gracillimus. Tubarao Series, Rio do Sul, S. Catarina, Bra- zil. "Ctenacanthus gradacostata St. John and COMMENTS: The type specimen is the Worth." (in) Caster (1930, p. 101). A typo- impression of a complete finspine in sand- graphical error for C. gradocostus (see below). stone. Its ornamentation is atypical of Cten- "Ctenacanthus gradocostatus St. J. and W." acanthus finspines and the species probably (in) Miller (1889) and Eastman (1907, p. 156). does not belong in this genus; it may be re- A typographical error for C. gradocostus (see ferable to Sphenacanthus. below). Ctenacanthus gracillimus Newberry and Ctenacanthus gradocostus St. John and Wor- Worthen (1866, p. 126, pl. XIII, fig. 3). then (1875, p. 425, pl. XV, figs. 2, 3). Plesiotype, Museum of Comparative Zo- Syntype, University of Illinois, X-358. ology, Harvard, MCZ 5184; Mississippian, Mississippian, Burlington Limestone, Quin- St. Louis Formation, St. Louis, Missouri; also cy, Illinois, Louisa Co., Indiana. from Alton, Illinois and Grand Rapids, COMMENTS: Said by St. John and Worthen Michigan. (1875) to "intimately" resemble C. burling- COMMENTS: The validity ofthis species has tonensis apart from minor differences in the been argued over since its inception. New- ornamentation. Although similar in general berry and Worthen (1866, p. 16, pl. XII, fig. shape to Acondylacanthus finspines, C. gra- 2) described another, much abraded finspine, docostus may represent a different, allied ge- as the type ofLeptacanthus occidentalis. This nus with more pectinate ornament. Another species was subsequently referred to Acon- specimen (AMNH 1034) bears a label in dylacanthus, and C. gracillimus was made Newberry's handwriting, noting similarities synonymous with A. occidentalis (St. John with "the type." and Worthen, 1875, p. 433). Later, however (St. John and Worthen, 1883, p. 238), it was Ctenacanthus gurleyi Newberry (1897, p. decided that preference should be given to 290). the name which was open to least doubt, and Mississippian, St. Louis Limestone, Salem, the name C. gracillimus was therefore used. Indiana (Gurley Coll.). Newberry (1889, p. 206) preferred to contin- COMMENTS: No type specimen is known, ue using the name A. occidentalis. Eastman nor was any specimen figured. These are suf- (1902, p. 85) followed St. John and Worthen ficient grounds to declare this species a no- (1883) in regarding gracillimus as the more men nudum, and it has elsewhere been sug- suitable specific name, but agreed with New- gested that it is synonymous with the type berry (1889) that this species should be re- species, Ctenacanthus major (Maisey, 1981). ferred to Acondylacanthus. Hussakof (1908) Ctenacanthus harrissi Caster (1930, p. 103, also used the name A. gracillimus. Zidek pl. 57, fig. 1). (1976) referred a finspine from the upper Mississippian, Mt. Jewett, Pensylvania. Mississippian of Oklahoma to Ctenacanthus COMMENTS: Caster (1930) suggests that C. aff. C. gracillimus. harrissi finspines resemble those of C. tenu- There is little doubt that C. gracillimus istriatus in the number of ribs and shape of should more correctly be referred to Acon- tubercles. This species is provisionally re- dylacanthus. The question of synonymy with tained in Ctenacanthus although it was not A. occidentalis cannot be resolved since that listed by Maisey (1981). 8 AMERICAN MUSEUM NOVITATES NO. 2774

Ctenacanthus harrisoni St. John and Wor- species to Sphenacanthus, though for very then (1883, p. 236, pl. XXIII, fig. 1). unsatisfactory reasons. Nevertheless, Maisey Type specimen, (?) USNM 13505, Missis- (1982) retains the species in Sphenacanthus sippian, St. Louis Limestone, Alton, Illinois. on other criteria. The type specimen of C. COMMENTS: See also Maisey (1981, fig. 6) marshi is very similar to S. serrulatus fin- where the species is retained in Ctenacanthus spines. as a possible synonym of C. major. Newberry (1889) noted similarities to C. littoni. See also Ctenacanthus ianishevskyi Khabakov (1928, Zidek (1977, p. 153, fig. I B). p. 23, pl. III, figs. 5-10). Holotype, CNIGR 2421/2, Leningrad; Ctenacanthus heterogyrus McCoy (1855, p. Mississippian (Carboniferous Limestone C), 625, pl. 3 I, fig. 32). Kuznetsk basin, Siberia. Mississippian, Carboniferous Limestone COMMENTS: Quite atypical for Ctenacan- Series, Armagh, Northern Ireland; Tournai, thus, the species has been transferred to By- Belgium. thiacanthus by Maisey (1982). The closest COMMENTS: First mentioned by Agassiz North American forms are C. solidus and C. (1837, vol. III, p. 177, name only), see also lucasi (also transferred to Bythiacanthus). DeKoninck (1878, p. 66, pl. VII, fig. 3), Davis (1883, p. 336, pl. XLIV, figs. 1-3), Wood- Ctenacanthusjaekeli Gross (1933, p. 64, pl. ward (1891, p. 101), Maisey (1982, p. 11, fig. II, fig. 9). 7). The type specimen of C. dubius was listed Type specimen, Geologisch-Palaontolo- under C. heterogyrus by Woodward (1891, gisches Museum, Berlin; Upper Devonian, p. 101). According to Khabakov (1928, p. 28, Wildungen. pl. IV, figs. 1-3) C. venator is similar to C. COMMENTS: Gross (1933) noted similari- heterogyrus; this suggestion was provision- ties between this form and Euphyacanthus ally accepted by Maisey (1982) pending ex- semistriatus Traquair (1894); this in turn re- amination of C. venator, but it seems possible sembles Tristychius in its finspine morphol- that C. venator is actually referable to Sphen- ogy (Dick, 1978). Referred to Homacanthus acanthus. Ctenacanthus heterogyrus was and regarded as an acanthodian by Denison transferred to Eunemacanthus by Maisey (1979, p. 52). The type specimen is unfor- (1982). tunately now lost, probably as a result of bombing in World War II (Dr. K. Fischer, Ctenacanthus hybodoides Egerton (1853, p. personal commun.). 280, pl. XII). Plesiotype, MCZ 5206 (formerly MCZ Ctenacanthus keokuk St. John and Worthen 4213), plesiotype from Mississippian coal (1875, p. 427, pl. XV, fig. 8). measures of Glasgow, Scotland, also wide- Type specimen (?), USNM 418, Mississip- spread in British Pennsylvanian coal mea- pian, Keokuk Limestone, Boonville, Mis- sures. souri. COMMENTS: See also Thomson (1874), COMMENTS: Supposed type specimen does Woodward (1889), Maisey (1982). The type not agree with figures, which may be com- specimen of C. nodosus was referred to this posites. Probably referable to Acondylacan- species by Woodward (1889) although they thus. are again separated by Maisey (1982). Thom- son (1869) incorrectly referred spines of this Ctenacanthus laevis Davis (1883, p. 341, pl. species to C. major (the type species), and XLV, fig. 1). reported an associated find of finspines with Type specimen, BM(NH) P2531, Missis- a partial dentition of Cladodus mirabilis (see sippian, Carboniferous Limestone, Armagh. also T. P. Barkas, 1873, pp. 19, 21; Thomson, COMMENTS: Woodward (1891, p. 102) sug- 1874). Subsequently W. J. Barkas (1874, gested that the species pertains to Acondyl- 1878) referred these finspines to Hybodus and acanthus, but the ornamentation and shape recommended suppression ofthe genus Cten- of their finspines disagree. Placed in a new acanthus. Woodward (1889) first referred the genus, Amelacanthus, by Maisey (1982). 1984 MAISEY: CTENACANTHUS AGASSIZ 9

Ctenacanthus lamborni Wells (1944, p. 65, rudis and Batacanthus baculiformis. While pl. 1, figs. 1-6). their ornamentation patterns are similar, Type specimen, Ohio State University however, the forms of the spines are rather Geological Museum No. 19501, upper Penn- different. The affinities of C. longinodosus are sylvanian, Ames Limestone, Guernsey Co., obscure, but it does not seem referable to Ohio. Ctenacanthus. COMMENTS: According to Wells (1944), at least two previously described specimens re- Ctenacanthus lucasi Eastman (1902, p. 80, ferred to C. amblyxiphias actually pertain to pl. 6, fig. 1, text-fig. 9). C. lamborni (in Eastman, 1903, and Branson, Type=specimen, USNM 4844, Mississip- 1916). A finspine like that of C. lamborni was pian, Kinderhook Limestone, Iowa. found associated with a toothplate of Me- COMMENTS: Another specimen, USNM gactenopetalus kaibabanus (see Hansen, 1978, 4686, is referred to this species. Maisey (1982) figs. 3-6). transfers the species to Bythiacanthus. Ctenacanthus latispinosus Whiteaves (1881, Ctenacanthus magnus (in) Thomson (1874, P. 99). p. 59). Lower Devonian, Campbellton, New COMMENTS: This is an invalid name. In an Brunswick. earlier publication, Thomson (1869, p. 102) COMMENTS: See also Whiteaves (1889, p. referred a specimen to Ctenacanthus major. 95, pl. X, fig. 3). Referred to the acanthodian The specimen was reassigned to C. hybo- genus Climatius by Woodward (1889, p. 183; doides by Thomson (1874), but in a footnote 1891, p. 33), and Denison (1979, p. 25). (p. 59) he incorrectly cited the previous iden- tification as C. magnus. Ctenacanthus mag- Ctenacanthus limaformis Davis (1883, p. 339, nus must therefore be regarded as an invalid pl. XLIV, fig. 5). junior synonym of Sphenacanthus hybo- Type specimen, BM(NH) P2535, Missis- doides. sippian, Carboniferous Limestone, Bristol, England. Ctenacanthus major Agassiz (1837, vol. III, COMMENTS: The type specimen is a badly p. 10, table 4). damaged finspine. The preserved ornament Type specimen, City of Bristol Museum is reminiscent of C. vetustus, but it is unlikely No. C4152, Mississippian, Carboniferous that either species is referable to Ctenacan- Limestone Series, Avon Gorge, Bristol (wide- thus. spread distribution recorded elsewhere). COMMENTS: The type species of Ctenacan- Ctenacanthus littoni Newberry (1889, p. 201, thus. See also Davis (1883, p. 334, plate XLII, pl. XXV, fig. 3). figs. 1, 2), Maisey (1981, figs. 1, lOL). Spec- Type specimen, AMNH 1050 (G), Missis- imens referred to this species by Trautschold sippian, St. Louis Limestone, St. Louis, Mis- (1874a, 1874b) are cephalopods (Khabakov, souri. 1928). Thomson (1869, p. 102) referred a COMMENTS: See also Hussakof (1908) and Sphenacanthus finspine to C. major (Wood- Maisey (1981), who retain the species in ward, 1889, p. 242), as did Newberry (1873; Ctenacanthus. Newberry (1889) noted sim- see Maisey, 1981, 1982). The following ilarity with C. harrisoni. species were regarded as synonyms of C. ma- Ctenacanthus longinodosus Eastman (1902, jorby Woodward (1891) and Maisey (1981): p. 78, text-fig. 8, pl. 5, fig. 2). C. tenuistriatus, C. maximus and C. salo- Paratypes, MCZ 5182 and USNM 3393, piensis. In addition, Maisey (1981) suggested Mississippian, "derived from the Kinder- that the following species may be synony- hook near mous with C. major: C. varians, C. varians Limestone," probably Burlington, var. russakovi, C. speciosus, C. spectabilis, C. Iowa ("North hill exposure"). and COMMENTS: Eastman (1902) noted a re- harrisoni, C. deflexus. semblance between the type specimen and Ctenacanthus maranhensis Silva Santos spines of Oracanthus, Glymmatacanthus (1946, p. 282, pl. 1, figs. 1, 2). 10 AMERICAN MUSEUM NOVITATES NO. 2774

Type specimen, DGM 448-P, National Triassic, middle Phosphoria Formation, Geological Museum, Brazil. Lower Permian, Wind River Mountains, Wyoming. Formacao Pedra do Fogo, Maranhao, Brazil. COMMENTS: This species is based on an in- COMMENTS: The species is based on a frag- determinate fragment of a finspine. The ment of finspine which, from its shape in "ridges" noted by Branson appear to be no transverse section and ornamentation pat- more than areas in between grooves for vas- tern, is not referable to Ctenacanthus. Instead cular canals. The only ornament is an enam- it may be closer to Sphenacanthus or to Wod- eled area on the presumed anterior margin. nika. This is reminiscent of Nemacanthus, and it Ctenacanthus marshi Newberry (1873, p. 326, may be that C. mutabilis is an early neose- pl. XXXVI, fig. 3). lachian. It is not referable to Ctenacanthus. Type specimen, YPM 2873, Pennsylva- Ctenacanthus nodocostatus Hussakof and nian, Coal Measures, Lanesville. Bryant (1918, p. 159, pl. 51, fig. 1). COMMENTS: Referred to Sphenacanthus by Type specimen, Buffalo Museum, E2083, Maisey (1982); the type specimen of C. mar- upper Devonian, Catskill Formation, "Sec- shi closely resembles S. hybodoides and S. ond Mountain Sandstone," Venango County, serrulatus finspines. Pennsylvania. Ctenacanthus maximus De Koninck (1878, COMMENTS: See also Maisey (1981, fig. 1OK) p. 68, pl. VII, fig. 1). where the species is retained in Ctenacan- Type specimen, IRSNB P1305, Mississip- thus. pian, Carboniferous Limestone, Soignies, Belgium. Ctenacanthus obscuracostatus Branson (1916, COMMENTS: Generally regarded as a syn- p. 654, pl. IV, fig. 2, text-figs. 2, 3). onym of C. major, the type species (Davis, Type specimen, University of Missouri, 1883; Woodward, 1891; Maisey, 1981). No.710, Permo-Carboniferous, lower Embar Formation, Big Popo Agie Canyon, Wyo- Ctenacanthus mayi Newberry and Worthen ming. (1870, p. 372, pl. II, fig. 2). COMMENTS: An indeterminate species based Type specimen, University of Illinois, on a finspine that resembles those of Acon- X- 1166, Mississippian, "lowerCarboniferous dylacanthus more than those of Ctenacan- Limestone," Burlington, Iowa. thus. COMMENTS: The ornament pattern and shape of the finspine in transverse section Ctenacanthus ornatus Agassiz (1837, p. 12, seem to preclude this species from Cten- pI. II, fig. 1). acanthus, but its affinities are obscure. Devonian, Old Red Sandstone. Another specimen, USNM 14184, represents COMMENTS: Generally considered to be the lower part of a similar finspine, with a based on a fragment of climatiiform acan- very deep transverse section, not unlike some thodian spine, and referred to Climatius; see Bythiacanthus finspines. also Woodward (1891, p. 32), Pageau (1969, Ctenacanthus minor Davis (1879b, p. 531, p. 455, fig. 16), Denison (1979, p. 25). with fig.). Ctenacanthus panderi (Eichwald): Wood- Pennsylvanian, lower Coal Measures ward (1889, p. 306). (Black-Bed Coal), near Bradford, Yorkshire, Mississippian, Carboniferous Limestone, England. Toula, U.S.S.R. COMMENTS: The type specimen is very small COMMENTS: Originally referred to Hybodus and does not seem to be fully formed. It may (Eichwald, 1860, p. 1603), the species is therefore pertain to a juvenile individual, "founded upon a spine probably of Ctena- probably of Sphenacanthus (Woodward, canthus" (Woodward, 1889, p. 306). This is 1889, p. 244; Maisey, 1982, p. 19). unhelpful, however, as Woodward's concept Ctenacanthus mutabilis Branson (1933, p. of Ctenacanthus spines was rather broad. In- 180, fig. 1, nos. 13, 14). terestingly, it was not subsequently included 1 984 MAISEY: CTENACANTHUS AGASSIZ I1I in Woodward's (1891) list of Ctenacanthus Ctenacanthuspugiunculus St. John and Wor- species. then (1875, p. 430, pl. XXI, fig. 9). Type specimen, USNM 13525, Mississip- Ctenacanthus parvulus Newberry (1875, p. pian, St. Louis Limestone, St. Louis, Mis- 55, pl. LIX, fig. 3). souri. Upper Devonian, Cleveland Shale, Bed- COMMENTS: The type specimen is a long, ford, Ohio. slender spine. It does not resemble the stout, COMMENTS: Newberry (1875) referred this heavily tuberculate spine referred to this species (which seems to be based on an im- species by Newberry (1897, p. 288, pl. XXII, mature spine) to Ctenacanthus with misgiv- fig. 4). His view that the species pertains to ings. It has subsequently been referred to Oracanthus seems untenable (see also Den- Hoplonchus (Newberry, 1889, p. 169; ison, 1979, p. 36). It is probably not referable Woodward, 1891, p. 107; Hussakof, 1908, p. to Ctenacanthus, however, but may be allied 49; Denison, 1979, p. 52). It is possible, how- to Asteroptychius. Another specimen sup- ever, that the species is founded on a juvenile posedly ofthis species (AMNH 5285) cannot Ctenacanthus finspine, since the tip ofa sim- be located at the time of writing. Ctenacan- ilar spine is present on AMNH 189 (the ar- thus similis is close to the present species and ticulated Ctenacanthus described by Dean, the two may be synonymous. 1909). Ctenacanthuspustulatus Davis (1883, p. 344, Ctenacanthuspellensis St. John and Worthen pl. XLV, fig. 2). (1883, p. 237, pl. XXI, fig. 2). Type specimen, British Museum (Natural Type specimen, USNM 13514, Mississip- History) BM(NH) P2529, Mississippian, pian, St. Louis Formation, Pella, Iowa. Lower Carboniferous Limestone, Armagh, COMMENTS: This species is founded on a Northern Ireland. small fragment of finspine which closely re- COMMENTS: Woodward (1891, p. 102) sug- sembles the type specimen of C. venustus. gested that the species may pertain to Astero- The two species may therefore be synony- ptychius. Maisey (1982) referred it to a new mous, and C. pellensis is retained in Cten- genus, Amelacanthus. acanthus here although it was not listed pre- Ctenacanthus randalli Newberry (1889, p. viously (Maisey, 1981). Another fragment was 105, no illustrations). described and figured by Branson (1906, p. Type specimen, AMNH 6675, Frasnian, 1393, fig. 25), from the Salem Limestone of Olean conglomerate (Chemung Group), near Salem, Indiana. Warren, Pennsylvania. Ctenacanthusperegrinus Khabakov (1928, p. COMMENTS: See also Eastman (1907, p. 25, pl. III, figs. 1-4). 154), Hussakof (1908, p. 46), Caster (1930, Holotype, CNIGR 2421/3, Leningrad, p. 102), and Maisey (1981, p. 20, fig. 8E). Carboniferous Limestone, Cl zone (Tour- Retained in Ctenacanthus by Maisey (1981). naisian), Roika Village, Tom River, Siberia. Ctenacanthus rectus Davis (1883, p. 345, pl. COMMENTS: The strong pustulose orna- XLV, fig. 5). ment of the figured specimen is reminiscent Mississippian, "Mountain Limestone," of Bythiacanthus; see Maisey (1982, fig. 3C- Armagh, Northern Ireland. F). COMMENTS: First listed by Agassiz (1843, Ctenacanthus plicatus Davis (1883, p. 342, vol. III, p. 177, name only) as Onchus rectus. pl. XLV, fig. 4). An indeterminate species. Mississippian, "Mountain Limestone," Ctenacanthus salopiensis Davis (1883, p. 339, Armagh, Northern Ireland. pl. XLIV, fig. 6). COMMENTS: First listed as Onchus plicatus Type specimen, BM(NH) P2523, Missis- (Agassiz, 1843, vol. III, p. 177, name only). sippian, Carboniferous Limestone, Oreton, Referred to a new genus, Amelacanthus, by Shropshire, England. Maisey (1982, fig. 5F-I). COMMENTS: A synonym of C. major, ac- 12 AMERICAN MUSEUM NOVITATES NO. 2774 cording to Woodward (1891, p. 98) and rulatus by Agassiz (1845) seem to pertain to Maisey (1981, p. 19). Psammosteus meandrinus Agassiz (1845) (see Ctenacanthus sculptus St. John and Worthen Woodward, 1891, p. 126), which has nothing (1875, p. 421, pl. XIV, fig. 1). to do with Sphenacanthus. See Agassiz (1837, Type specimen, USNM 13520, Mississip- vol. III, p. 24, pl. I, figs. 11-13) and Maisey pian, upper Kinderhook Beds, lower (no. 1) (1982, fig. 8A). fish bed, Burlington, Iowa. Ctenacanthus similis St. John and Worthen COMMENTS: Probably close to Ctenacan- (1875, p. 431, pl. XV, fig. 11). thus, but ornament pattern is not pectinate. Syntype, University of Illinois, X- 1157, Nonetheless, the tubercles on the costae are Mississippian, Chester Limestone, Chester, arranged transversely. Tubercles become Illinois. coarser anteriorly, though less strongly than COMMENTS: This species is close to C. pugi- in C. vetustus, with which C. sculptus may be unculus and they may be synonymous. These allied. Both species may eventually be re- elongate slender spines are not referable to moved from the genus Ctenacanthus. Another Ctenacanthus, but have yet to be assigned to specimen, USNM 6048, from St. Louis, Mis- a different genus. souri, is referred to C. sculptus. Ctenacanthus solidus Eastman (1902, p. 90, Ctenacanthus semicostatus (St. John and pl. 7, fig. 3, text-fig. 13). Worthen): Eastman (1902, p. 89). Type specimen, USNM 3383, paratypes, Type specimen, MCZ 5187, Mississippian, USNM 4843, MCZ 5185, Mississippian, upper Burlington Group, Burlington, Iowa. Kinderhook Formation, Iowa and Illinois. COMMENTS: The genoholotype of Anaclit- COMMENTS: Eastman (1902, pl. 7, fig. 3) acanthus St. John and Worthen (1875, p. 443, figured the paratype, USNM 4843, only; see pl. XVI, fig. 14). The type specimen is not a Maisey (1982, fig. 3A) for the type. Eastman's Ctenacanthus finspine, and Eastman (1902) text-figure 7 is misleading as the spine is much is hardly justified in his remarks that its "re- deeper than he suggests anteroposteriorly, but lations are evidently with the genus Ctena- is damaged so that its posterior wall has bro- canthus instead of Anaclitacanthus." Most ken away. The species is transferred from probably the type specimen is a paired spine, Ctenacanthus to Bythiacanthus by Maisey possibly acanthodian. There are interesting (1982). Paratype MCZ 5185 is recognizable similarities with C. cylindricus. The pectoral as pertaining to this species, but has suffered spines of Lupopsyrus pygmaeus Bernacsek from deformational shattering within the and Dineley (1977), a climatiid acanthodian, matrix. are similar to the type specimen of Anacli- Ctenacanthus speciosus St. John and Wor- tacanthus, but have fewer and coarser ridges. then (1875, p. 424, pl. XIV, figs. 3, 4). Also similar is Acondylacanthus? mugdianus Type specimen, USNM 13518, Mississip- of St. John and Worthen (1883, p. 244, pl. pian, Kinderhook Formation, "lower fish XXIV, fig. 3). bed," Iowa. Ctenacanthus serrulatus (Agassiz): Traquair COMMENTS: Synonymous with C. varians (1884, p. 6) non-Agassiz (1845, p. 119. pl. according to Eastman (1902, p. 89). Also may XXXIII, fig. 24). be synonymous with C. major according to Type specimen, RSM. GY. 1878.18.15. Maisey (1981, p. 19). Calciferous sandstones (Mississippian), Bur- Ctenacanthus spectabilis St. John and Wor- diehouse, near Edinburgh, Scotland. then (1875, p. 420, pl. XV, fig. 1). COMMENTS: This is the type species of Type specimen, USNM 13519, Mississip- Sphenacanthus Agassiz (1837, vol. III, p. 24), pian, Kinderhook Beds, La Grande, Iowa. which Traquair (1884) regarded as a syn- COMMENTS: See also Eastman (1902, p. 87, onym of Ctenacanthus, although elsewhere pl. 5, fig. 1) and Maisey (1981, fig. 5). these genera are considered distinct (e.g., Woodward, 1889, p. 242; Maisey, 1981, Ctenacanthus sulcatus (Agassiz): Davis (1883, 1982). Spines referred to Ctenacanthus ser- p. 343, pl. XLV, fig. 3). 1984 MAISEY: CTENACANTHUS AGASSIZ 13

Type specimen, Bristol City Museum, ical of Ctenacanthus, but is like that of Wod- C4154, Mississippian, "Mountain Lime- nika. stone," Armagh, Northern Ireland. COMMENTS: The species was first described Ctenacanthus varians St. John and Worthen (as Onchus sulcatus) by Agassiz (1837, vol. (1875, p. 422, pl. XIV, fig. 2). III, p. 8, table 1, fig. 6). Included in Cten- Type specimen, MCZ 5186, Mississippian, acanthus by Davis (1883) and Woodward Kinderhook Series, "upper fish bed," Flint (1891), but made the type species of a new River, near Burlington, Iowa. genus, COMMENTS: See also Eastman (1902, p. 89), Amelacanthus, by Maisey (1982). Maisey (1981, fig. 4). See C. speciosus, which Ctenacanthus tenuirostris von Meyer (1854, is probably synonymous. The present species p. 53, pl. VIII, figs. 18, 19, 20). may be synonymous with C. major (Maisey, COMMENTS: An inadequately described 1981, p. 19). species, regarded by De Koninck as identical Ctenacanthus varians St. John and Worthen with C. maximus. Apparently von Meyer was var. russakovi Khabakov (1928, p. 26, pl. IV, referring to C. tenuistriatus (see below) but figs. 11-14). got the name wrong (see also Woodward, Holotype, CNIGR 2421/1, Leningrad, 1891, p. 98). This is an awkward reference Lower Carboniferous, Cl zone, Kirghizian to retrieve, since Woodward (1891) incor- Steppes, Siberia. rectly lists it as "Roemer, 1850." Not only COMMENTS: May be synonymous with C. was the volume published in 1854 (Roemer's major (Maisey, 1981, p. 19). contribution being delivered in 1850), but von Meyer actually described the fish remains in Ctenacanthus near C. varians St. John and that work (pp. 53-54). Von Meyer suggests Worthen: Dorr and Moser (1964, p. 108, figs. that some teeth resembling those of "Dicre- 1-5). nodus" or "Carcharopsis" may have per- Mid-Mississippian, Michigan Formation, tained to C. tenuistratus, which is founded Grand Rapids, Michigan: UMMP 45738. upon a fragmentary spine having Ctenacan- COMMENTS: Seems to resemble the type thus-like ornamentation. specimen quite closely. Associated with cop- rolite pellets. Ctenacanthus tenuistriatus Agassiz (1837, vol. III, p. 11, table 3, figs. 7-11). Ctenacanthus venator Khabakov (1928, p. 28, Paratypes, BM(NH) P495, 2225, Bristol pl. IV, figs. 1-3). City Museum (no catalogue number, now Mississippian, C1-C2 zones, Donetz Basin, lost). Mississippian, "Mountain Limestone," Lissichia Balka. Bristol. COMMENTS: Said by Khabakov (1928) to COMMENTS: See also De Koninck (1878, p. resemble C. heterogyrus, the figured speci- 67, pl. VII, fig. 2), Davis (1883, p. 335, pl. men bears a clear resemblance to Sphen- XLIII, figs. 1, 2), Maisey (1982, fig. 3). Re- acanthus; see also Maisey (1982, fig. 7J-M). garded as a synonym of C. major by Wood- The holotype is apparently lost (O. Lebedev, ward (1891, p. 98) and Maisey (1981, p. 19). personal commun., May 1982). Ctenacanthus triangularis Newberry (1873, Ctenacanthus venustus Eastman (1902, p. 81, p. 329, pl. XXXVI, fig. 1). pl. 3, fig. 2, text-fig. 10). Type (?) specimen (cast), AMNH 423 (G), Holotype, MCZ 5183, plesiotype USNM Mississippian, Waverly Series, Oil Creek, 3385, Mississippian, Kinderhook Group, Pennsylvania. ?Iowa. COMMENTS: The wax cast AMNH 423 (G) COMMENTS: Eastman (1902) figured the agrees with Newberry's (1873) published fig- plesiotype. This and the holotype are figured ure. The whereabouts of the natural mould by Maisey (1981, fig. 1 OA-J). Other referred is unknown. Said to have been collected in specimens are USNM 4684 and USNM 3381 association with teeth like those of Orodus (two specimens). The species was retained in and Psammodus. Finspine ornament is atyp- Ctenacanthus by Maisey (1981). 14 AMERICAN MUSEUM NOVITATES NO. 2774

Ctenacanthus vetustus Newberry (1873, p. C. acutus; probably not referable to Cten- 326, pl. XXXV, fig. 3). acanthus. Type specimen, AMNH 351, Upper De- Ctenacanthus sp. (Hussakofand Bryant, 1918, vonian, Cleveland Shale, Sheffield Town- p. 161), Buffalo Museum no. E2498, De- ship, Lorain Co., Ohio. vonian, Genesee Series, Conodont Bed, Erie COMMENTS: See also Newberry (1889, pl. Co., New York; may be a distinct species. XXVIII, fig. 5), Hussakof (1908, p. 46). Fin- Ctenacanthus sp. (Branson and Mehl, 1938, spines associated with Orodus variabilis teeth p. 123, pl. 37, fig. 20). University of Mis- in CMNH 8103A, 8103B, and assigned to a souri no. 730 VP, lower Mississippian, new, but as yet undescribed, genus (Hlavin, Chouteau Province, Missouri. An indeter- 1972, 1976). minate fragment of spine, probably not referable to Ctenacanthus. No ornament is Ctenacanthus wrightii Newberry (1884, p. preserved. 206, pl. XVI, figs. 12-14). Ctenacanthus sp. (Kulczycki, 1957, p. 285, Type specimen, AMNH 352, Middle De- pl. XIII, fig. 2), Famennian, lower Chei- vonian (Erian), Moscow Shale, Hamilton loceras Beds, Kielce, Holy Cross Moun- Group, Yates Co., New York. tains, Poland. Similar to Acondylacanthus COMMENTS: See also Newberry (1889, p. and resembles C. gracillimus except for de- 66, pl. XXVI, fig. 4), Eastman (1907, p. 153), tails of ornamentation pattern. Hussakof( 908, p. 46), Hussakofand Bryant Ctenacanthus sp. indet. (Dorr and Moser, (1918, p. 161, pl. 52, fig. 2). Spelt wrighti by 1964, pp. 110 and 111, pl. 1, figs. 6, 8). all but Newberry (1884, 1889). These specimens (UMMP 23845 and Ctenacanthus? xiphias St. John and Worthen 45739) are indeterminate impressions of (1883, p. 244, pl. XVI, fig. 1). spines. Their ornamentation is atypical of Type (?) specimen, USNM 3391, Missis- Ctenacanthus. sippian, Keokuk Limestone, Keokuk Rapids, Ctenacanthus sp. (Bendix-Almgreen and near Keokuk, Iowa. Malzahn, 1969, p. 44; see also Schaum- COMMENTS: Founded on an abraded and berg, 1977, p. 308, fig. 9), may be a neo- worn specimen, this indeterminate species selachian. Schaumberg (1982) has referred was referred tentatively to Acondylacanthus this specimen to a new genus and species, by St. John and Worthen (1883). In shape Hopleacanthus richelsdorfensis, and sev- and overall form, however, the type (?) spec- eral articulated specimens are now known imen seems more like a Ctenacanthus fin- from the Permian Kupferschiefer of Ger- spine (see also Eastman, 1902, p. 86). many. "Ctenacanthus type" (Bendix-Almgreen, 1975, p. 551), Upper Carboniferous, Greenland. REFERENCES TO Ctenacanthus, Ctenacanthus sp. (Janvier, 1977, p. 131, fig. SPECIES INDETERMINATE 1C), Upper Devonian, Iran (Iran National Museum ofNatural History, MMTT 5005). The following references are arranged "Indeterminate ctenacanthid" spines (Jan- chronologically. vier, 1977, p. 129, fig. IA, B); Devonian, Ctenacanthus sp. (Davis, 1876, p. 335, no. Aroma Province, Brazil. 4), probably Sphenacanthus. Ctenacanthus sp. (Zidek, 1977, p. 151, fig. Ctenacanthus sp. (Davis, 1876, p. 335, no. 1A), Pennsylvanian, Seminole Formation, 5), probably Sphenacanthus. Tulsa County, Oklahoma. Said to have Ctenacanthus sp. (Eastman, 1902, p. 76), similarities with C. buttersi, but may be MCZ 5189, Pennsylvanian, Carlinville, Il- referable to Sphenacanthus. linois; probably Sphenacanthus. Ctenacanthus sp. (Zidek, 1977, p. 151, fig. Ctenacanthus sp. indet. (Eastman, 1902, p. 1 B), Pennsylvanian, Coffeyville Forma- 83), USNM 3480, Mississippian, Keokuk tion, Tulsa County, Oklahoma. Said to have Limestone, near Keokuk, Iowa. Close to similarities with C. harrisoni. 1984 MAISEY: CTENACANTHUS AGASSIZ 15

DISCUSSION ably does not belong to this genus, and I con- The following species of Ctenacanthus sider this to be an acanthodian species. Cten- should be considered invalid: acanthus longinodosus may also be an acanthodian. buttlersi (misspelling of buttersi) The following species have been, or prob- crenulatus (name only-later described as ably should be, referred to another form-ge- crenatus) nus, Acondylacanthus: fallax (indeterminable) gradacostata (misspelling of gradocostus) browni gradocostatus (misspelling of gradocostus) burlingtonensis gurleyi (no holotype or figure) cannaliratus magnus (incorrect citation) distans rectus (indeterminable) gemmatus tenuirostris (misspelling of tenuistriatus) gracillimus wrighti (misspelling of wrightil) keokuk obscurocostatus The type specimens ofthe following species The following species have been, or prob- are known to be lost or destroyed and no ably should be, referred to the genus Astero- neotypes have been designated: ptychius: crenatus bellus elegans pugiunculus gemiindensis simi/is jaekeli venator The following species have been, or probably should be, referred to Sphenacanthus: The following species are left as indeterminate, either because they are founded on aequistriatus inadequate material, or because the original costellatus descriptions are vague and imprecise, or be- depressus cause the material has not been examined gondwanus and its whereabouts not yet established: hybodoides magnus (invalid-see above) limaformis (badly damaged holotype) marshi panderi (inadequate description) minor (juvenile) parvulus (fragment of spine apex) serrulatus xiphias (damaged holotype-no details visi- ble) The following species have been, or prob- The following species have been referred ably should be, referred to other genera: to acanthodians: acutus (resembles Tristychius) abnormis (to Machaeracanthus) bosnensis (to Nemacanthus) bohemicus (to Machaeracanthus) brevis (to Bythiacanthus) cylindricus (resembles buttersi (hybodont?) Gyracanthus) costatus (to Eunemacanthus) erectus ("Onchus"?) dubius (to Amelacanthus?) gemundensis (to Nodacosta: holotype de- excavatus (to Eunemacanthus?) stroyed) gradocostus (close to Acondylacanthus) latispinosus (to Climatius) heterogyrus (to Eunemacanthus) ornatus (to Climatius) ianishevskyi (to Bythiacanthus) semicostatus (Homalacanthus; resembles laevis (to Amelacanthus) Lupopsyrus) lucasi (to Bythiacanthus) Homacanthus deliculatus was referred to maranhensis (close to Sphenacanthus or Ctenacanthus by Denison (1979), but it prob- Wodnika) 16 AMERICAN MUSEUM NOVITATES NO. 2774 mayi (resembles Bythiacanthus) species thought to be synonymous with C. mutabilis (resembles Nemacanthus) major are discounted) is as follows: peregrinus (resembles Bythiacanthus) Lower Carboniferous (Mississippian)- plicatus (to Amelacanthus) eight species. pustulatus (to Amelacanthus) Upper Devonian-Seven species. This is a solidus (to Bythiacanthus) relatively restricted geological range, whereas triangularis (hybodont?) "Ctenacanthus" sensu lato (i.e., all the species listed here) ranges from the Lower Devonian The following list of species still referred (e.g., C. abnormis, C. bohemicus, C. latispi- to Ctenacanthus is slightly revised from nosus, all coincidentally now referred to Maisey (1981): acanthodians) to the Triassic (e.g., C. mu- angulatus tabilis, C. bosnensis). All the species retained angustus in Ctenacanthus are from marine deposits. chemungensis Only two of these species (the late Devonian clarkii C. clarkii and C. compressus) are known from cliftonensis articulated remains (Maisey, 1981). compressus deflexus (synonym of C. major) denticulatus ACKNOWLEDGMENTS formosus I acknowledge with many thanks the help harrissi I have received from numerous people harrisoni (synonym of C. major) worldwide in digging up obscure references, littoni holotype numbers, and historical facts about major (type species) disappearances and reappearances of speci- maximus (synonym of C. major) mens. Any mistakes in the information re- nodocostatus main mine, however, and I will be glad to pellensis receive further information concerning the randalli species listed here as well as others that I may salopiensis (synonym of C. major) have inadvertently overlooked. speciosus (synonym of C. major) spectabilis (synonym of C. major) LITERATURE CITED tenuistriatus (synonym of C. major) Agassiz, L. varians (synonym of C. major) 1833-1844. Recherches sur les poissons fos- venustus siles. Neuchatel, 5 vols., 1420 pp. The following species are founded in fin- 1844-1845. Monographie des poissons fossiles that resemble those of Ctenacanthus du vieux gres rouge ou systeme Devo- spines nien (Old Red Sandstone), des Iles Bri- but are sufficiently different (in my view) to tanniques et de Russie. Neuchatel, xxxvi preclude them from that genus: + 171 pp. amblyxiphias (has continuous pectinate keel Barkas, T. P. anteriorly) 1873. Illustrated guide to the fish, amphibian, reptilian and supposed mammalian re- coxianus (atypical ornament) mains of the Northumberland Carbon- decussatus (atypical ornament) iferous strata. London, Hutchings, viii furcicarinatus (atypical ornament) + 117 pp. lamborni (has continuous pectinate keel an- Barkas, W. J. teriorly) 1874. Hybodus, a Coal Measure fish. Geol. sculptus (atypical ornament) Mag., n.s., vol. 1, pp. 163-168,239,287- vetustus (atypical ornament) 288. wrightii (atypical ornament) 1878. Ctenacanthus, a spine ofHybodus. Jour. & Proc. Roy. Soc. New South Wales, vol. l1,pp. 145-155. The stratigraphic distribution ofthe species Barrande, J. retained in the genus Ctenacanthus (when all 1872. Systeme Silurien du centre de la Bo- 1984 MAISEY: CTENACANTHUS AGASSIZ 17

heme. 1. Partie: Recherches paleonto- fishes. Proc. U.S. Natl. Mus., vol. 14, logiques. Supplement au vol. 1. Trilo- pp. 447-463. bites, crustaces divers et poissons. Davis, J. W. Prague, xxx + 647 pp. 1876. On a bone-bed in the lower Coal Mea- Bendix-Almgreen, S. E. sures, with an enumeration of the fish- 1975. The palaeovertebrate faunas of Green- remains of which it is principally com- land. In Escher, A., and W. S. Watt posed. Quart. Jour. Geol. Soc., vol. 32, (eds.), Geology of Greenland. Copen- pp. 332-340. hagen, Geol. Surv. Greenland, pp. 536- 1879a. Notes on Pleurodus affinis Agassiz, and 573. descriptions of three spines of cestra- Bendix-Almgreen, S. E., and E. Malzahn - - cionts from the lower Coal Measures. 1969. Ueber neue oder wenig bekannte Elas- Ibid., vol. 35, pp. 181-188. mobranchier aus dem deutschen Kup- 1879b. Description of a new species of fossil ferschiefer. Hessisches Landesamt Bo- fish spine, Ctenacanthus minor, from the denforsch., vol. 97, pp. 44-45. lower Coal Measures ofYorkshire. Geol. Berman, D. S. Mag., vol. 6, pp. 531-532. 1970. Vertebrate fossils from the Lueders For- 1883. On the fossil fishes ofthe Carboniferous mation, lower Permian of north-central Limestone series ofGreat Britain. Trans. Texas. California Univ. Publ. Geol. Sci., Roy. Soc. Dublin, vol. 1, pp. 327-600. vol. 86, 39 pp. Dean, B. Bernacsek, G. M., and D. L. Dineley 1909. Studies on fossil fishes (sharks, chimae- 1977. New acanthodians from the Delorme roids and arthrodires). Mem. Amer. Formation (lower Devonian) ofN.W.T., Mus. Nat. Hist., vol. 9, pp. 209-287. Canada. Palaeontographica, Abt. A, vol. De Koninck, L. G. 158, pp. 1-25. 1878. Faune de Calcaire Carbonifiere de la Branson, C. C. Belgique. Ann. Mus. Roy. d'Hist. Nat. 1933. Fish fauna of the middle Phosphoria de Belgique, vol. 2, pp. 1-150. Formation. Jour. Geol., vol. 41, no. 2, Denison, R. pp. 174-183. 1979. Handbook of paleoichthyology. Vol. 5. Branson, E. B. Acanthodii (Schultze, H.-P., ed.). New 1906. Fish remains from the Salem Limestone York, Gustav Fischer Verlag, vi + 62 of Indiana. 30th Ann. Rept., Depart. PP. Geol. & Nat. Res., Indiana, pp. 1376- Dick, J. R. F. 1394. 1978. On the Carboniferous shark Tristychius 1916. The lower Embar of Wyoming and its arcuatus Agassiz from Scotland. Trans. fauna. Jour. Geol., vol. 24, pp. 639-664. Roy. Soc. Edinburgh, vol. 70, pp. 63- 1933. Kinderhookian of Missouri (abstract). 109. Proc. Geol. Soc. 352. America, p. Dorr, J. A., and F. Moser Branson, E. B., and M. G. Mehl 1964. Ctenacanth sharks from the mid-Mis- 1938. Pisces from the lower Mississippian of sissippian of Michigan. Papers Michi- Missouri. In Branson, E. B., M. G. Mehl, gan Acad. Sci., vol. 49, pp. 105-113. A. K. Miller, R. Peck, I. Keyte, and W. Furnish (eds.), Stratigraphy and paleon- Eastman, C. R. tology of the lower Mississippian of 1897. On Ctenacanthus spines from the Keo- Missouri, II. Univ. Missouri Studies, kuk limestone ofIowa. Amer. Jour. Sci., vol. 13, no. 4, pp. 109-127. vol. 3, pp. 10-12. 1902. Some Carboniferous cestraciont and Caster, K. E. acanthodian sharks. Bull. Mus. Comp. 1930. Higher fossil faunas of the upper Alle- Zool., vol. 39, pp. 55-99. ghenys. Bull. Amer. Paleont., vol. 15, 1903. Carboniferous fishes from the central no. 58, pp. 1-175. western states. Ibid., vol. 39, pp. 163- Claypole, E. W. 226. 1885. On Ctenacanthus and Gyracanthus from 1907. Devonian fishes of the New York for- the Chemung of Pennsylvania. Proc. mations. Mem. N.Y. State Mus., vol. Amer. Assoc. Adv. Sci., 1884, pp. 489- 10, pp. 1-235. 490. Egerton, P. M. G. Cope, E. D. 1853. On two new species of placoid fishes 1891. On the characters of some Paleozoic (Ctenacanthus hybodoides and Ct. no- 18 AMERICAN MUSEUM NOVITATES NO. 2774

dosus) from the Coal Measures. Quart. Hussakof, L., and W. L. Bryant Jour. Geol. Soc., vol. 9, pp. 280-282. 1918. Catalogue of the fossil fishes in the mu- Eichwald, C. E. von seum of the Buffalo Society of Natural 1853-1861. Lethaea Rossica, ou paleontologie Sciences. Buffalo Soc. Nat. Sci., Bull. 12, de la Russie decrite et figuree. Stuttgart, 346 pp. vol. 1, pp. 1493-1607. Janvier, P Fritsch, A. 1977. Description des restes d'elasmobranches 1895. Fauna der Gaskohle und der Kalksteine (Pisces) du Devonien Moyen de Bolivie. der Permaformation B6hmens. Prague, Palaeovertebrata, vol. 7, part 4, pp. 126- vol. 3, 132 pp. 132. Giebel, C. G. 1858. Die silurische Fauna des Unterharzes Katzer, F. nach Herrn C. Bischof's Sammlung 1916. Prilozi paleontologiji Bosne i Hercego- bearbeitet. Abhandl. Naturwiss. Ver., vine. 1. Ctenacanthus bosnensis n. f. Je- Provinz Sachsen und Thuringen, vol. 1, dan novi Ichthyodorylith iz trijasa. pp. 263-332. Glasnik Zemaljskog Museja u Bosni i Gillette, D. D., and E. A. Shapiro Hercegnovini, vol. 28, nos. 1-2, pp. 201- 1977. Catalogue of type specimens of fossil 206. vertebrates, Academy of Natural Sci- Kayser, F. H. E. ences, Philadelphia; Part V, Fishes. Proc. 1878. Die Fauna der altesten Devon-Ablage- Acad. Nat. Sci. Philadelphia, vol. 129, rungen des Harzes. Abhandl. Geol. Spe- pp.113-123. cialk. Preussen, vol. 2, no. 4, 296 pp. Gross, W. Khabakov, A. V. 1933. Die Wirbeltiere des Rheinischen De- 1928. Description of new species of ichthyo- vons. Abhandl. Preussischen Geol. dorulites ofthe genus Ctenacanthus Ag. Landesanst., Heft 154, p. 83. from Carboniferous deposits in the 1940. Acanthodier und Placodermen aus den USSR. Izvestiya Geol. Komiteta, vol. Heterosteus-Schichten Estlands und 47, pp. 23-31 (in Russian with English Lettlands. Ann. Soc. Reb. Nat. Invest. summary). Univ. Tartuensi Const., vol. 46, pp. 12- Koenen, A. 99. 1895. Ueber einige Fischreste des nord- Hansen, M. C. deutschen und b6hmischen Devons. 1978. A presumed lower dentition and a spine Abhandl. Gesellsch. G6ttingen, vol. 40, of a Permian petalodontiform chon- no. 2, pp. 1-37. drichthyan, Megactenopetalus kaibabanus. Jour. Paleont., vol. 52, no. 1, pp. Kulczycki, J. 55-60. 1957. Upper Devonian fishes from the Holy Hlavin, W. J. Cross Mountains (Poland). Acta Pale- 1972. New associations of fossil sharks from ont. Polonica, vol. 2, pp. 285-382. the Cleveland Shale, Upper Devonian Leidy, J. (Famennian). Geol. Soc. America, Abst. 1857. Descriptions of the remains of fishes with programs, vol. 4, p. 21. from the Carboniferous Limestone of 1976. Biostratigraphy of the late Devonian Illinois and Missouri. Trans. Amer. Phil. black shales on the cratonal margin of Soc., vol. 2, pp. 87-90. the Appalachian geosyncline. Ph.D. Maisey, J. G. Dissertation, Boston Univ. Graduate 1981. Studies on the Paleozoic selachian ge- School, 1976, Geology, 194 pp. (Uni- nus Ctenacanthus Agassiz. No. 1. His- versity Microfilms Internatl., 76- torical review and revised diagnosis of 21,285.) Ctenacanthus, with a list ofreferred taxa. Hussakof, L. Amer. Mus. Novitates, no. 2718, pp. 1- 1908. Catalogue of the type and figured spec- 22. imens offossil vertebrates in the Amer- 1982. Studies on the Paleozoic selachian ge- ican Museum of Natural History. Part nus Ctenacanthus Agassiz. No. 2. By- I. Fishes. Bull. Amer. Mus. Nat. Hist., thiacanthus St. John and Worthen, vol. 25, pp. 1-103. Amelacanthus, new genus, Eunema- 1911. The Permian fishes of North America. canthus St. John and Worthen, Sphena- Carnegie Inst. Washington Publ. No. canthus Agassiz, and Wodnika Mun- 146, pp. 153-178. ster. Ibid., no. 2722, pp. 1-24. 1984 MAISEY: CTENACANTHUS AGASSIZ 19

McCoy, F. Pageau, Y. 1848. On some new fossil fish of the Carbon- 1969. Nouvelle faune ichthyologique du De- iferous period. Ann. Mag. Nat. Hist., vonien moyen dans le gres de Gaspe ser. 2, vol. 2, pp. 1-10, 115-133. (Quebec). II. Morphologie et systema- 1855. Descriptions of the British Palaeozoic tique. Premiere Section. A. Eurypte- fossils in the Geological Museum ofthe rides. B. Ostracodermes. C. Acantho- University of Cambridge. Cambridge diens et Selachiens. Nat. Can., vol. 96, Univ. Press, 661 pp. pp. 399-478. Meyer, H. von Romer, A. S. 1854. Squaliden-Reste aus dem Posidono- 1942. Notes on certain American Paleozoic myen Schiefer des Oberharzes bei Ober- fishes. Amer. Jour. Sci., vol. 240, pp. Schulenberg, pp. 53-54. In Roemer, F. 216-228. Beitrage zur geologischen Kenntniss des Schaumberg, G. nordwestlichen Harzgebirges. Palaeon- 1977. Der Richelsdorfer Kupferschiefer und tographica, vol. 3, pp. 1-67. seine Fossilien. III. Die tierischen Fos- Miller, S. A. silien des Kupferschiefers. 2. Vertebra- 1889. North American geology and paleon- ten. Der Aufschluss, vol. 28, pp. 297- tology for the use of amateurs, students 352. and scientists. Cincinnati, Press of 1982. Hopleacanthus richelsdorfensis n.g. n. Western Methodist Book Concern, 664 sp., ein Euselachier aus dem permischen PP. Kupferschiefer von Hessen (W- Moy-Thomas, J. A. Deutschland). Palaont. Z., vol. 56, pp. 1936. The structure and affinities of the fossil 235-257. elasmobranch fishes from the lower Silva Santos, R. Carboniferous rocks of Glencartholme, 1946. Duas novas formas de Elasmobran- Eskdale. Proc. Zool. Soc. London (1936), quios do Paleozoico de Meio do Norte, pp. 761-788. Brasil. An Acad. Brasil Cien., vol. 18, Newberry, J. S. no. 4, pp. 282-284. 1873. Description of fossil fishes. Ohio Geol. 1947. Um Ctenacanthus do Gondwana Bra- Surv., vol. 1, part 2, pp. 245-255. siliero. Ibid., vol. 19, no. 3, pp. 248- 1875. Description of fossil fishes. Ibid., vol. 2, 250. part 2, pp. 1-64. 1878. Description of new Palaeozoic fishes. St. John, 0. H., and A. H. Worthen Ann. N.Y. Acad. Sci., vol. 1, pp. 188- 1875. Description of fossil fishes. Geol. Surv. 192. Illinois, Geol., Paleont., vol. 6, pp. 245- 1884. Ctenacanthus wrighti, n. sp. 35th Ann. 488. Rep. N.Y. State Mus. Nat. Hist., 1884, 1883. Description of fossil fishes. Ibid., vol. 7, p. 206. pp. 55-264. 1889. The Palaeozoic fishes of North Ameri- Thomson, J. ca. U.S. Geol. Surv. Monograph XVI, 1869. On teeth and dermal structures associ- 340 pp. ated with Ctenacanthus. Rep. Brit. As- 1897. A new species and a new genus ofAmer- soc. Adv. Sci., 39th meeting, 1869, pp. ican Palaeozoic fishes, together with 102-103. notes on the genera Oracanthus, Dac- 1874. On Ctenacanthus (Sphenacanthus) hy- tylodus, Polyrhizodus, Sandalodus, Del- bodoides Egerton. Trans. Geol. Soc. todus. Trans. N.Y. Acad. Sci., vol. 16, Glasgow, vol. 4, pp. 59-62. pp. 282-304. Traquair, R. H. Newberry, J. S., and A. H. Worthen 1884. Description ofa fossil shark (Ctenacan- 1866. Description of vertebrates. Geol. Surv. thus costellatus) from the lower Car- Illinois, vol. 2, Palaeontology, pp. 9- boniferous rocks of Eskdale, Dumfries- 141. shire. Geol. Mag., vol. 1, pp. 3-8. 1870. Description of fossil vertebrates. Ibid., 1894. Notes on Palaeozoic fishes. No. 1. Ann. vol. 4, pp. 347-374. Mag. Nat. Hist., vol. 14, pp. 368-374. Obruchev, D. V. Trautschold, H. von 1967. Fundamentals ofpaleontology. Vol. 1, 1874a. Fischreste aus dem Devonischen des Agnatha, Pisces. Israel Program for Sci- Gouvernements Tula. Nouv. Mem. Soc. entific Translations, U.S. Dept. ofCom- Imp. Nat. Moscou, vol. 13, pp. 263- merce, x + 825 pp. 276. 20 AMERICAN MUSEUM NOVITATES NO. 2774

1874b. Die Kalkbriiche von Mjatschkova. Eine Roy. Soc. Canada, vol. 6, sec. 4, pp. 77- Monographie des oberen Bergkalkes. 96. Ibid., vol. 13, pp. 277-326. Woodruff, E. G. Tuomey, M. 1906. The geology of Cass County, Nebraska. 1858. Descriptions and figures of Ctenacan- Geol. Surv. Nebraska, vol. 2, pp. 169- thus elegans, Cladodus newmani, C. 292. magnificus. In Mallet, J. W. (ed.), Sec- Woodward, A. S. ond Biennial Report Geol. Alabama, pp. 1889. Catalogue ofthe fossil fishes in the Brit- 38-40. ish Museum (Nat. Hist.), Part I. Lon- Wells, J. W. don, British Mus. (Nat. Hist.), xliv + 1944. A new fish spine from the Pennsylva- 474 pp. nian of Ohio. Ohio Jour. Sci., vol. 44, 1891. Catalogue ofthe fossil fishes in the Brit- pp. 65-67. ish Museum (Nat. Hist.), Part II. Ibid., Whiteaves, J. F. xliv + 567 pp. 1881. On some fossil fishes, crustacea and Zidek, J. mollusca from the Devonian rocks at 1976. Oklahoma paleoichthyology. Part V. Campbellton, N. B., with descriptions Chondrichthyes. Oklahoma Geol. of five new species. Canadian Natural- Notes, vol. 36, no. 5, pp. 175-192. ist, vol. 10, pp. 93-101. 1977. Oklahoma paleoichthyology. Adden- 1889. Illustrations of the fossil fishes of the dum to Part V. Ibid., vol. 37, no. 5, pp. Devonian of Canada. Part IL. Trans. 151-156.