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Permian Sharks of Kansas:

In 2003, Keith Ewell discovered an unusual occurrence of Permian shark remains in Kansas

Copyright © 2003-2011 by Mike Everhart

Updated 09/09/2011




Left: Assorted Cladodus teeth from Keith Ewell's Permian shark tooth site #1. 

The remains of Permian sharks (290-248 mya) are not unknown from Kansas but are certainly less well known than those from the underlying Pennsylvanian (323-290 mya) rocks. However, it appears that sharks were not evolving rapidly during this 75 million year period and most remains are identifiable from Pennsylvanian species that continue on into the Permian.

Note that the identification of cladodont sharks is somewhat confused at this point.  Most species are based on isolated teeth or dorsal fin spines, not on associated remains.  According to Maisey (pers. comm, 2003), Petalodus teeth and the curved fin spines called Physonemus may be from the same shark. Others have suggested Cladodus teeth and Ctenacanthus dorsal fin spines may be from the same genus. In any case, I am interested in hearing any suggestions on the identifications we have made so far.

Ewell, K. and Everhart, M.J. 2005. A Paleozoic shark fauna from the Council Grove Group (Lower Permian). Kansas Academy of Science, Transactions 108(1-2):71-72 (Abstract).

In 2003, abundant vertebrate remains consisting of unassociated shark teeth, dorsal fin spines and preserved cartilage were discovered in a narrow (2-3 cm) seam of poorly consolidated limey mud near the base of the Neva Limestone Member of the Genola Limestone Formation (Council Grove Group, Lower Permian) in southeastern Geary County, Kansas. Remains identified included the teeth of the Paleozoic sharks Cladodus occidentalis, Petalodus alleghaniensis, Agassizodus variabilis, Acrodus sp., Chomodus sp., and a chimaeroid (cf.  Sandalodus sp.), dorsal fin spines of Physonemus mirabilis and a hybodont shark (cf. Lissodus sp.), and scales of Holmesella quadrata. The mud layer contains a high percentage of coarsely ground shell and bone fragments, and is interpreted as a re-deposit generated by a major storm or tsunami type event. Another nearby site produced dorsal fin spines of Ctenacanthus cf. amblyxiphias, teeth of Cladodus sp. and Petalodus sp., and preserved cartilage in association with abundant invertebrate remains, while teeth of Petalodus sp. and Chomodus sp., and an unidentified dorsal fin spine fragment were collected at a third site. Cladodus teeth were the most frequently recovered remains at Site 1, while Ctenacanthus dorsal fin spines and Petalodus teeth were the most common remains at Site 2 and Site 3 respectively. Differing conditions of deposition are indicated by the condition and abundance of vertebrate remains at each site.

Site 1 - Unless otherwise indicated, all of the teeth shown below were collected by Keith Ewell, and all come from the same shark-tooth-rich horizon in the Council Grove Group (Lower Permian), in western Wabaunsee County, Kansas. The site has produced six or more species of shark remains, including teeth, dorsal fin spines and calcified cartilage from a relatively small exposure.  There may be as many as 30 or 40 teeth per square meter in this layer. Most of the specimens shown below are curated in the collections of the Sternberg Museum of Natural History, Hays, Kansas.

ke-005a.jpg (14182 bytes) Left: Large Cladodus occidentalis Leidy 1859 tooth in matrix, labial view. While these teeth are often complete, the are also usually fractured.  (KE-005)

Right:  Smaller Cladodus occidentalis tooth (views: upper row, left to right - lingual, labial, and occlusal; bottom row, left to right - lingual, lateral, lateral, and basal. (KE-013)

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ke-043a.jpg (10754 bytes) Left: Lingual and labial views of a very large Cladodus sp. tooth (Base width = 36 mm). The crown and accessory cusps on this tooth exhibited a lot of natural wear (KE-043).

Right: Crown and basal views of a slightly larger Cladodus sp. tooth (Base width = 38 mm) - (KE-042).

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ke-014a.jpg (11643 bytes) Left: A nice Cladodus occidentalis tooth (KE-014). Click here for similar teeth in the collection of the University of Kansas (KUVP 65013).

Right: Large "Petalodus alleghaniensis"  (A) lingual view; B) labial view; C) occlusal view; D) enlarged labial view - note "serrations" along cutting edge. Only a few Petalodus teeth have been found at Site #1 (KE-016)

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ke-022a.jpg (7076 bytes) Left: Smaller teeth, possibly Orodus ? (KE-022)

Right: Acrodus ? ((KE-017)

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ke-023a.jpg (7895 bytes) Left: Orodus ? (KE-023)

Right: Two Acrodus? teeth (KE-024)

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ke-018a.jpg (8975 bytes) Left: Small cladodont tooth identified as "Maiseyodus johnsoni" (unpublished species, Hansen, 1986). (KE-018). (Click here for a scan of Hansen's Figure 18 <WI>

Right: A small fragment from the upper end of a  Ctenacanthus dorsal fin spine.   Note the characteristic denticles on the posterior edge of the spine. (KE-026)

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vp24001a.jpg (8261 bytes) Left: A very large dorsal fin spine of Physonemus mirabilis in right lateral view in the University of Kansas collection (Upper Pennsylvanian - See Chorn and Frailey, 1978). The insert shows the detail of the posterior denticles. (KUVP 25001)

Right: A nearly complete dorsal fin spine of Physonemus mirabilis (Xystracanthus arcuatus Leidy) in right lateral view. Posterior denticles HERE. (KE-015)

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ke-039a.jpg (8459 bytes) Left: An unusual spine with sharp, teeth-like posterior denticles, the tips of which are broken in this specimen. Possibly from the hybodont shark, Lissodus. (KE-039) <WI>

Right: A second specimen of the spine of the same organ genus exhibiting sharp posterior denticles. (KE-040)

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ke-021a.jpg (5505 bytes) Left: These teeth are probably from Agassizodus variabilis (corrugatus), a "Cestraciont" shark first described from Kansas by Professor B. F. Mudge in 1874.  (KE-021) <MG>

Right: As above (KE-029)

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ke-038a.jpg (8762 bytes) Left: These teeth are probably also from Agassizodus, a "Cestraciont" shark first described from Kansas by Professor B. F. Mudge in 1874.  (KE-038) <MG>

Right: As above (KE-030)

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ke-037a.jpg (8668 bytes) Left: These teeth are probably from Agassizodus, a "Cestraciont" shark first described from Kansas by Professor B. F. Mudge in 1874.  (KE-037) <MG>

Right: As above (KE-032)

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agassiza.jpg (19424 bytes) Left:  Detail from Portion of Plate VIII (St. John and Worthen, 1875) showing the Agassizodus corrugatus jaw plate found in Osage County, Kansas.

Right: Larger portion of Plate VIII (St. John, and Worthen, 1875) showing the Agassizodus corrugatus jaw plate from Osage County, Kansas.

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vp55215a.jpg (14199 bytes) Left and Right: Two views of 4 teeth of Campodus (Agassizodus) corrugatus teeth in natural articulation from University of Kansas collection (KUVP 55215). Found by R. E. Fox in 1910 in the "Coal measures."


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campod1a.jpg (9209 bytes) A second, articulated specimen of "Campodus" found in the Pennsylvanian of Jefferson County, KS. (KU collection, found by J. Savage) campod3a.jpg (9604 bytes)campod2a.jpg (8166 bytes)
ke-041a.jpg (11731 bytes) Left: A complete Chomodus sp. tooth (KE-041).

Right: Unknown tooth fragment, probably Chomodus. (KE-033)

Far Right: Unknown tooth  fragment (KE-025)

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Left: Sandalodus (Deltodus)? Tooth crown fragment. Note that this tooth is from a chimaeroid fish. Chimaeras, or Ratfishes are cartilaginous fishes, but are not sharks. (KE-020)

Right: Detail of calcified cartilage. (KE-034)


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ke-010a.jpg (9355 bytes) Left: Holmesella quadrata scale. (KE-010) <DC>

Right: Two Holmesella quadrata scales. This genus was previously reported by Ørvig (1966) from Upper Pennsylvanian rocks in Montgomery Co., Kansas)    (KE-019)

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Site 2 -Geary County #1 -This is the second of Keith's Permian shark sites, somewhat higher in the section than Site 1. Keith has found a number of Ctenacanthus spines at this site, two of which occurred next to a Cladodus tooth (probably more of a coincidence than an association). While spines are plentiful, teeth are rare.

2ke-003a.jpg (16585 bytes) Left: A Ctenacanthus amblyxiphias Cope 1891 dorsal fin spine in the matrix where it was found by Keith. This is the second nearly complete spine of this species found in Kansas. (2KE-003)

Right: The anterior end of the fin spine, complete to the rounded tip. (2KE-005)

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2ke-004a.jpg (14946 bytes) Left: The possible association of this Ctenacanthus amblyxiphias fin spine and a broken Cladodus tooth. There were only two teeth were found in this exposure; both were Cladodus and both were found within a foot (30 cm) of fin spines. (2KE-004)

Right: Detail of area where this fin spine inserted into the body of the shark. (2KE-006)

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2ke-008a.jpg (7924 bytes) Left: Detail of lateral ornamentation on the anterior portion of the fin spine, near the insertion border. Details of ridge on the anterior edge HERE.

Right: Note similarity in this detail of the lateral ornamentation on the anterior portion of the dorsal fin spine of Ctenacanthus amblyxiphias (FHSM VP-15012)

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2ke-010a.jpg (10211 bytes) Left: Nearly complete dorsal fin spine of Ctenacanthus amblyxiphias Cope. One of three fin spines found by KE on 09/11/2003.

Right: Large dorsal fin spine of Ctenacanthus sp.

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2ke-007a.jpg (11119 bytes) Left: Occlusal view of the Cladodus tooth, missing the central cusp, found close to the fin spine above. (2KE-007)

Right: A large piece of calcified cartilage, possibly from the braincase or lower jaws of a shark, or the base of the dorsal fin.  Unfortunately, nothing else was found. (2KE-003)

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ke-027a.jpg (12045 bytes) Left: A second Ctenacanthus amblyxiphias dorsal fin spine fragment that I found within a foot of the tooth at right: (KE-027)

Right: An occlusal view of a Cladodus tooth, missing the central cusp, found by Keith Ewell. (KE-028)

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2ke-012a.jpg (11278 bytes) Left: A very small Cladodus tooth. (2KE-012)

Right: An unknown cladodont tooth, probably not Cladodus. (2KE-013)

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Site 3 - Geary County #2: This is another of Keith's sites, below Site 2 but somewhat higher in the section than Site 1 (still lower Permian). This site has numerous Petalodus teeth, but few teeth of other species.

3ke-002a.jpg (9538 bytes) Left: The results of an hour of hunting at Keith's Site 3. Lots of Petalodus teeth, and not much else. See larger version of the low-crowned, posterior tooth (arrow) below left. (3KE-002)

Right: Two large Petalodus teeth: Top complete crown, missing root; Bottom, Chipped crown, nearly complete root. (3KE-001)

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3ke-004a.jpg (8767 bytes) Left: Results of the second day of hunting at the same site.  Three of the four fragments are from one tooth. (3KE-004)

Right: Large and nearly complete Petalodus tooth.

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3ke-003a.jpg (6859 bytes) Left: Nearly complete crown of a posterior tooth of Petalodus. (3KE-003)

Right: A very complete, posterior tooth of Petalodus.

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3ke-009a.jpg (7767 bytes) Left: Chomatodus arcuatus St. John; partial tooth. (3KE-009)

Right: A very large Petalodus tooth found by Keith Ewell's mother, Kathy (10/2003) at site 3. Tooth measures 5 cm by 5 cm. (3KE-010)

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3ke-006a.jpg (7599 bytes) Left: Fragment of a ctenacanth spine. This may be from a shark called Amelacanthus as figured in Maisey (1982). (3KE-006) <WI>






Chorn, J. and D. Frailey. 1978. A large chondrichthyan spine, Physonemus mirabilis, from the Upper Pennsylvanian of Kansas, U.S.A. N. Jb. Geol. Paläont. Mh., (7):385-392, 1 fig.

Cicimurri, D. 2002.  Chondrichthyes from the upper part of the Minnelusa Formation (Middle Pennsylvanian: Desmoinesian), Meade County, South Dakota. Proceedings of the South Dakota Academy of Science, 81:81-92.

Cope, E.D. 1891. Characters of Paleozoic fishes. Proc. U.S. Nat. Museum. 14:447-463. (Ctenacanthus amblyxiphias Cope 1891)

Everhart, M.J. and P. A. Everhart. 2003. First report of the Paleozoic shark, Ctenacanthus amblyxiphias Cope 1891 from the Lower Permian of Morris County Kansas. Kansas Acad. of Science (Abstracts), 22:13.

Ewell, K. and Everhart, M.J. 2005. A Paleozoic shark fauna from the Council Grove Group (Lower Permian). Kansas Academy of Science, Transactions 108(1-2):71-72 (Abstract).

Hussakof, L. 1911. The Permian Fishes of North America, In Case, E. C., Revision of the Amphibia and Pisces of the Permian of North America.  Carnegie Institute of Washington. (Ctenacanthus amblyxiphias Cope 1891)

Itano, W. M., K. J. Houck and M. G. Lockley. 2003. Ctenacanthus and other chondrichthyan spines and denticles from the Minturn Formation (Pennsylvanian) of Colorado. Journ. Paleon. 77(3):524-535.

Leidy, J. 1859. [Xystracanthus, Cladodus and Petalodus from the Carboniferous of Kansas]. Proc. Acad. Nat. Sci. Phila., 11:3.

Maisey, J.G. 1975. The interrelationships of phalacanthous selachians. N. Jb. Geol. Paläont. Mh. 9:553-567, 6 fig.

Maisey, J.G. 1977. Structural notes on a cladoselachian dorsal spine. N. Jb. Geol. Paläont. Mh. 1:47-55, 6 fig.

Maisey, J.G. 1982. Studies on the Paleozoic selachian genus Ctenacanthus No. 2. Bythiacanthus St. John and Worthen, Amelacanthus St. John and Worthen, Sphenacanthus Agassiz, and Wodnika Münster. American Museum Noviates, 2722:1-24.

Maisey, J.G. 1984. Studies on the Paleozoic selachian genus Ctenacanthus Agassiz. No. 3.: Nominal species referred to Ctenacanthus. American Museum Noviates, 2774:1-20.

Mazzullo, S.J., C.S. Teal, and C. A. Burtnett. 1997. Outcrop stratigraphy and depositional facies of the Chase Group (Permian, Wolfcampian) in Kansas and southeastern Nebraska. Kansas Geological Survey Technical Series 6, 210 p.

Mudge, B. F. 1874. Rare forms of fish in Kansas. Kansas Acad. Sci. Trans. 3:121-122 (Agassizodus tooth plate)

Ørvig, T. 1966. Histological studies of Ostracoderms, Placoderms, and fossil Elasmobranchs. 2. On the dermal skeleton of two late Palaeozoic Elasmobranchs. Ark. Zool. Stockholm. 19(1):1-39.

Robb, A.J. 2003. Notes on the occurrence of some petalodont shark fossils from the Upper Pennsylvanian rocks of northeastern Kansas. Kansas Academy of Science, Transactions 106(1-2):71-80.

Romer, A.S. 1942. Notes on certain American Paleozoic fishes. American Journal of Science, 240:216-228, 1 pl. 

Schultze, H-P. 1985. Marine to onshore vertebrates in the Lower Permian of Kansas and their paleoenvironmental implications. University of Kansas , Paleontological Contributions, Paper 113:1-18.

Schultze, H.-P., J. D. Stewart, A. M. Neuner and R. W. Coldiron, 1982. Type and figured specimens of fossil vertebrates in the collection of the University of Kansas Museum of Natural History.  Part I. Fossil fishes. Misc. Pub. Univ. Kansas Mus. Nat. Hist. 73:53 pp. (see p. 5 for a listing of Holmesella sp. scales from the Pennsylvanian of Kansas) <EM>

St. John, O.A. and H. Worthen. 1875. Palæontology: Descriptions of fossil fishes. Geology and Palæontology, Part 2, Section 1, Geological Survey of Illinois, 6:245-532, i-vi, pl. 1-22.

Zangerl, R. 1981. Chondrichthyes I - Paleozoic Elasmobranchii. 3a:1-115 In Shultze, H-P. (ed.), Handbook of Paleoichthyology. Gustav Fischer Verlag, Stuttgart and New York.

Zidek, J. 1976. Oklahoma Paleoichthyology Part V: Chondrichthyes. Oklahoma Paleontology Notes, 36(5):175-192, Oklahoma Geol. Surv., Norman, OK.  

Credits: I thank Keith Ewell for the loan of these specimens and the opportunity to learn more about Paleozoic sharks.  David Cicimurri (Clemson University), Michal Ginter (University of Warsaw, Poland), Mike Hansen (Ohio Geological Survey),  Wayne Itano ( NIST, Boulder, CO), John Maisey (American Museum of Natural History), Earl Manning (Tulane University), Bill May (Sam Noble Museum, Norman, OK), and Kenshu Shimada (DePaul University) have assisted in the identifications and/or provided reference material.