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A Field Guide to Fossils of the Smoky Hill Chalk


Part 2:

Sharks and Bony Fish (Continued from Part 1: Invertebrates)


Copyright © 2000-2013  by Mike Everhart


Latest update 07/22/2013



CLICK TO ENLARGE --- "We're Not in Kansas Any More. Copyright © Ray Troll; used with permission of Ray Troll.   Ray's website is here: TROLLART.COM


VERTEBRATES: Animals having a spinal column, including fish, reptiles and birds. Fossils are generally limited to skeletal remains but sometimes also include calcified cartilage, scales and stomach contents. (My list of fish references is here


You can now download a copy of this early article on Kansas Sharks by Williston - Provided by the Kansas Geological Survey:

Williston, S. W. 1900. Cretaceous fishes: Selachians and Pycnodonts. University Geological Survey Kansas VI pp. 237-256, with pls.


Click here for Mark Wilson's Mesozoic Fishes Page  --- A list of references

Class: Chondrichthyes Huxley, 1880


Cartilaginous fish, including sharks:  


Sharks are an old family of fish, dating back many millions of years to the Devonian Period. They were common predators and scavengers in the Western Interior Sea. For picture of the teeth of more species, click here for Sharks of Kansas


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Sharks shed their teeth continuously through their life.  These teeth are the most commonly found vertebrate fossils.  Isolated teeth are frequently collected in the Smoky Hill Chalk.


LEFT: (CLICK FOR LARGER PICTURE) The most common kinds of shark teeth found in the Smoky Hill Chalk Member (Late Cretaceous) are: 1A) Squalicorax falcatus, 1B) Squalicorax kaupi;   2) Cretoxyrhina mantelli; 3) Cretalamna appendiculata;, and 4A) Ptychodus mortoni; 4B) Ptychodus martini, 4C) Ptychodus anonymus.


Cretoxyrhina mantelli:

"Ginsu" sharks grew lengths of 20 feet or more, and were one of the top predators in the Western Interior Sea. They were probably very similar in behavior to the modern Great White Shark, but were not closely related. Their razor sharp, triangular, blade-like teeth are the largest shark teeth found in the chalk and their large, disk shaped vertebrae are also found occasionally. Sometimes the entire shark including skin is preserved as a fossil. Charles Sternberg found a very large Cretoxyrhina in the late 1800s. On page 113-114 of C. H. Sternberg's "Life of a Fossil Hunter," he describes the discovery of a 20 [foot?] shark (Cretoxyrhina) from the chalk (probably Gove Co., south of Park, KS). The specimen was nearly complete, with "over 250 teeth"....

Quoting Sternberg, "This is the first time and, I believe, the only time that so complete a specimen of this ancient shark has been discovered. The
column and other solid parts were composed of cartilaginous matter which usually decays so easily that is rarely petrified. I suppose my specimen was old at the time of its death, and bony matter had  been deposited in the cartilage. It is not very likely that such a specimen will ever be
duplicated.  Dr. Eastman's [1895?] study of this skeleton enabled him to make synonyms of many species which had been named from teeth alone."

According to Sternberg, "the most complete skeleton of the Cretaceous shark, Oxyrhina mantelli [Oxyrhina = Cretoxyrhina] Agassiz ever discovered in any formation" was sold to the "Ludwig-Maximilian University of Munich." Apparently, the study he was referring to is: Eastman, C. R., 1895. Bietrage Kenntniss Gattung Oxyrhina mit besonderer Berucksichtigung von Oxyrhina mantelli Agassiz.  Palaeontographica 41:149-192.

Unfortunately, the specimen was destroyed during World War II.

LEFT: A drawing of a well preserved specimen of Cretoxyrhina mantelli (FHSM VP-323) at the Sternberg Museum in Hays, Kansas. (Drawing by Kenshu Shimada)


Read the story of the recent (2002) discovery of a giant, and very complete specimen of Cretoxyrhina mantelli.


LEFT: Calcified vertebral centra from a large Cretoxyrhina mantelli (FHSM VP-2184).  The specimen included over 100 centra, but only a few teeth.

RIGHT: Overview of the entire FHSM VP-2184 specimen. 


For some reason, Cretoxyrhina mantelli seems to reach it's largest size and greatest abundance in the lower one third of the chalk (Late Coniacian), and then, judging from the size of the shed teeth that are found, becomes rarer and smaller in the upper chalk. Cretoxyrhina then became extinct worldwide by the middle of the Campanian (82 mya).

cretox4a.jpg (11446 bytes) LEFT: Teeth of the ginsu shark,  Cretoxyrhina mantelli Agassiz (FHSM VP-2187).

RIGHT: Examples of the different shapes of Cretoxyrhina mantelli  teeth. The shape of the tooth indicates it's position in the jaw of the shark, with taller straighter teeth in the front and smaller teeth in the back of the jaws..

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Cretalamna appendiculata:

c-appn1a.jpg (11951 bytes) LEFT: A smaller shark than Cretoxyrhina, Cretalamna appendiculata has long, double curved teeth, with smaller, triangular cusplets on either side of the main shaft.  

RIGHT: Another lateral Cretalamna appendiculata tooth from the lower chalk (Late Coniacian) of Gove County, KS.  Relatively rare in the chalk, this species is more common in the lower chalk, and pre-Niobrara deposits.

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Scapanorhynchus r. raphiodon: Goblin Sharks

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LEFT: A group of five, associated Scapanorhynchus cf. S. raphiodon teeth found in southeastern Gove County, KS (Late Coniacian). These tiny teeth were found by Shawn Hamm.  They are quite rare in the Smoky Hill Chalk. (FHSM VP-13961)


RIGHT: A group of six, unassociated Scapanorhynchus teeth collected by Keith Ewell and myself in September, 2004, from the lower chalk of Trego County, KS.

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Johnlongia sp.

ke-shc1a.jpg (8060 bytes) LEFT: A tooth of Johnlongia sp. (FHSM VP-15545) from the Smoky Hill Chalk (Late Coniacian) of Trego County, KS. Found by Keith Ewell, 11/2003. This tooth represents the first record of this species in Kansas.

See: Shimada, K., K. Ewell and M. J. Everhart. 2004. The first record of the lamniform shark genus, Johnlongia, from the Niobrara Chalk (Upper Cretaceous, western Kansas. Kansas Academy of Science, Transactions 107(3/4):131-135.

RIGHT: Another Johnlongia sp. tooth collected in 2010 from Smoky Hill Chalk (Late Coniacian) of Trego County, KS. Scale = mm


Squalicorax falcatus / kaupi / pristodontus: Crow sharks

NOTE: While I can honestly take credit for being the first to use the term "Ginsu shark" as a common (but appropriate) name for Cretoxyrhina mantelli (above), the origin of the term "Crow Shark is a bit more mysterious. I don't know when it was first used or who used it, but I suspect it goes back to the meaning of the Greek word "Corax." The genus name, "Squalicorax" was originally just "Corax." (e.g. Corax falcatus Agassiz 1843). Corax is a name found in Greek mythology, but also is the species name for the common raven (a bird) of Europe (Corvus corax).  (From Wikipedia, in regard to Corvus corax, "the specific epithet, corax/???a?, is the Greek word for "raven" or "crow". Whitley (1939) changed the genus name to Squalicorax. At some point, Squalicorax was given the common name, Crow shark.

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These pictures of one of the most complete specimens of Squalicorax falcatus (USNM 425665) that has ever been found (United States National Museum - The Smithsonian). It was discovered in the Smoky Hill Chalk of western Kansas. See Shimada and Cicimurri (2005) for a complete description.

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In 1992, I was also fortunate enough to discover the front half of a 2 meter Squalicorax falcatus (below). The specimen (CMC VP 5722) was donated to the Cincinnati Museum Center. 


cmc5722w.jpg (7880 bytes) cmc5722x.jpg (9967 bytes) The specimen preserves the teeth roughly in the jaws... and the dermal denticles (scales, HERE and HERE) in the shark skin preserve the rough outline of the head. The preserved vertebral centra (calcified cartilage) are shown HERE.  See Shimada and Cicimurri (2005) for a description of this specimen. cmc5722y.jpg (11910 bytes) cmc5722z.jpg (9087 bytes)

Shimada, K. and Cicimurri, D.J. 2005. Skeletal anatomy of the Late Cretaceous shark, Squalicorax (Neoselachii: Anacoracidae). Paläontologische Zeitschrift 79(2): 241-261.

squali1a.jpg (13714 bytes) The genus Squalicorax evolved through several "species" during the deposition of the chalk in the Western Interior Seaway, approx. 87-82 million years ago. Squalicorax falcatus is known from the Middle Cenomanian in Kansas (See the earliest Kansas teeth from the middle Cenomanian Upper Dakota below).  S. kaupi begins to appear in small numbers during the early Santonian and pretty much replaces S. falcatus in Kansas by the early Campanian. A few S. pristodontus teeth have been found right at the top of the Smoky Hill Chalk. 
s-falc1a.jpg (15131 bytes) LEFT: Lingual and labial views of a typical Squalicorax falcatus tooth; From the middle Santonian Smoky Hill Chalk, Little Pyramids, Logan Co., Kansas.

RIGHT: Lingual and labial views of a posterior lateral tooth of Squalicorax kaupi, upper chalk (Lower Campanian), Logan County, Kansas.

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pscoraxa.jpg (22957 bytes) Left: Pseudocorax laevis (FHSM VP-15823) is a rare shark species found in the lower (Late Coniacian) Chalk. This tooth was found by Keith Ewell in 2004 in Trego County (below MU 2). Note the lack of serrations.

Right: An unusual  Squalicorax sp. tooth in labial (left) and lingual views that was found in situ under the recently discovered dinosaur vertebrae. Note the very weak serrations on the main cusp and the lack of serrations on the posterior accessory cusp.   (Upper Coniacian)

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S-kaupi2a.jpg (25674 bytes) Left: A Squalicorax kaupi tooth in lingual (far left) and labial views. Tooth was found in the Smoky Hill Chalk of Gove County above Hattin's marker unit 8.

Right: A typical Squalicorax kaupi tooth (FHSM VP-17566), from the Early Campanian chalk of Logan County. Note the change in the shape of the cusp and the root.

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VP2213-Ba.jpg (32162 bytes) Left: A portion of the preserved jaw cartilage of a Squalicorax kaupi in the Sternberg Museum (FHSM VP-2213) with 5 teeth in their natural arrangement (labial view).

Right: Detail of the jaw of FHSM VP-2213 showing two teeth in labial view.

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s-pris1a.jpg (13702 bytes) Left: A typical Squalicorax pristodontus tooth, but a rare discovery at the very top of the Smoky Hill Chalk - early Campanian, Logan Co, KS. This tooth and two others were collected by Pete Bussen.

Right: Squalicorax "what?" ... One of several specimens of an unusual tooth form from the lower Smoky Hill Chalk collected by Keith Ewell.

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Ptychodontids - Pavement-toothed or Crusher sharks

Family Ptychodontidae Jaekel, 1898

    Genus Ptychodus Agassiz, 1835


Ptychodontids: Possibly a ray-like shark, this genus had it's upper and lower jaws covered with rows of rounded, peg-like teeth that were used to crush the hard shells of small clams and oysters. It is likely that the Ptychodontid sharks fed by ingesting large amounts of bottom mud and sieving out the edible shellfish and other organisms. The teeth are usually found one at a time (scattered remains) but occasionally entire jaw plates with their many rows of teeth are found intact.



RIGHT: An artist's reconstruction of a ptychodontid shark in an mural at the Sternberg Museum of Natural History. Unfortunately, we have no complete specimens that would tell us if this form is correct or not.

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Four species (Ptychodus mortoni, P. martini and P. anonymus, and more rarely, P. polygyrus (also possibly P. latissimus) are found in the lower chalk. Ptychodontids become extinct in the Western Interior Sea and none are found above the middle of the chalk.


Leidy (1868) was the first to report on Ptychodus teeth from the "Cretaceous series" of western Kansas when he described a single, damaged tooth from "a few miles east of Fort Hays, Kansas." The tooth was obtained by Joseph LeConte during the survey for the Kansas Pacific Railroad in 1867. Leidy considered it to be a new species and named it Ptychodus occidentalis. Although the tooth was later lost from the collection of the Academy of Natural Sciences in Philadelphia (Spamer, et al., 1995), the figure of it shown in Leidy (1873, pl. 17 and 18) appears instead to be a marginal tooth of P. mortoni. Leidy (1873, pl. 18) also illustrates several P. mortoni teeth collected by Dr. George M. Sternberg while on military duty in western Kansas during the Indian wars of the late 1860s.


RIGHT: A composite (upper and lower teeth)  jaw plate (KUVP 55270) of Ptychodus mortoni in the University of Kansas Natural History Museum. The jaw plate is approximately 10 inches wide by 18 inches in length and includes more than 500 teeth. A close-up view is shown HERE. Note the wear on the tops of the teeth in the center rows. Anterior (labial) is to the bottom of the picture. This articulated specimen was collected by Prof. E. S. Rose in the vicinity of Castle Rock in Trego County. For more information about Ptychodus, see this very good, early paper:


Woodward, A. S., 1887. On the dentition and affinities of the selachian genus Ptychodus Agassiz. Quarterly Journal of the Geological Society xliii pp. 121-131, with pl. x.


... and:


Everhart, M. J. and Caggiano, T.  2004.  An associated dentition and calcified vertebral centra of the Late Cretaceous elasmobranch, Ptychodus anonymus Williston 1900. Paludicola 4(4),  p. 125-136.


However, note that Ptychodus anonymous Williston 1900 in the Smoky Hill Chalk has more recently been re-identified as Ptychodus rugosus Dixon 1850 (Hamm, 2010a), and P. polygyrus has been re-identified as P. marginalis (Hamm, 2010b):


Hamm, S.A. 2010a. The Late Cretaceous shark, Ptychodus rugosus (Ptychodontidae) in the Western Interior Sea. Kansas Academy of Science, Transactions 113(1-2):44-55.


Hamm , S.A. 2010b. The Late Cretaceous shark Ptychodus marginalis in the Western Interior Seaway, USA. Journal of Paleontology 84(3):538–548.

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olson1a.jpg (24431 bytes) Ptychodus is a genus of durophagous (shell-crushing) sharks from the Late Cretaceous. Their teeth are often found as fossils around the world, but most often in the sediments deposited in the Western Interior Sea of North America. They became extinct during the Santonian, about 85 million years ago.

LEFT and RIGHT: Associated upper and lower jaw plates of Ptychodus mortoni collected from middle Smoky Hill Chalk Gove Co., Kansas, and prepared by Dennis Olson.

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pt-ano1a.jpg (10074 bytes) Ptychodus anonymus Williston 1900 (now P. rugosus per Hamm, 2011) is one of 4 species of ptychodontid sharks found in the Smoky Hill Chalk.

Left: Ptychodus anonymus teeth, from the medial row.

Right: Ptychodus anonymus posterior lateral teeth (FHSM VP-14854) from the lower chalk of Gove County.

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pt-mor3a.jpg (10353 bytes) Ptychodus mortoni is the only ptychodontid species which has ridges radiating from the center of the tooth. All other species have ridges that run more or less parallel across the crown.

Left: Ptychodus anonymus (FHSM VP-2238) jaw plate in the collection of the Sternberg Museum. 

Right: Ptychodus mortoni (FHSM VP-336)

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pt-mor1a.jpg (11734 bytes) Left: Ptychodus mortoni teeth (FHSM VP- 335); Note the excessive wear on the tooth at far left.

Right: Detail of a Ptychodus mortoni lower jaw plate (FHSM VP-14785) that I found in Gove County in 1990. This specimen is fully described in Shimada (2012):

Shimada, K. 2012. Dentition of Late Cretaceous shark, Ptychodus mortoni (Elasmobranchii, Ptychodontidae). Journal of Vertebrate Paleontology 32:6:1271-1284.

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pt-mar1a.jpg (14856 bytes) Left: Ptychodus martini - 100+ associated teeth, Gove Co., KS (Late Coniacian). Found by Shawn Hamm in Gove County.

Right: Detail from the type specimen of Ptychodus martini Williston 1900 in the collection of the University of Kansas (KUVP 55271).

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pmartina.jpg (10361 bytes) Left: A badly weathered P. martini Williston 1900 found by Pam Everhart in the lower chalk of Gove County, KS. Note the flat aspect of the crown in both of these specimens. No other tooth crown of a Ptychodus species is quite this flat.

Right: Ptychodus martini -Left: A single tooth from the Late Coniacian Smoky Hill Chalk of Trego County, KS. Found by Keith Ewell, 11/2003.

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p_martia.jpg (17612 bytes) LEFT: Occlusal and medial (?) views of a tooth of Ptychodus martini found by Pam Everhart in the low chalk of Trego County (10/2004).  The identification is based on the description provided by S.W. Williston, 1898. 

RIGHT:  One of the smaller 'symphysial' teeth that serve as spacers between the two medial rows of large teeth in the upper jaw of a Ptychodus, probably P. mortoni (found by Keith Ewell, 10/2004).

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pt-pol1a.jpg (14114 bytes) Left: A large Ptychodus polygyrus (KUVP 55237) in the collection of the University of Kansas, This specimen was the subject of one of the first photographs ever published of shark teeth (Williston, 1900).

Right: Another large P. polygyrus (FHSM VP-76) at the Sternberg Museum.

Note that this species has now been re-identified by Hamm (2010)  in the Western Interior Seaway as Ptychodus marginalis.

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pt-pol2a.jpg (9432 bytes) Left: A Ptychodus polygyrus in the collection of the Sternberg Museum (FHSM VP-15008); Gove Co., KS (Late Coniacian). Found by Tom Caggiano.

Right: Ptychodus mammillaris lateral tooth -FHSM VP-14022, from near the base of the Fort Hays Limestone, (Lower Coniacian) , Ellis Co., KS. Collected by Mike Everhart.

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Rhinobatos incertus - Guitar fish

shcrhina.jpg (13995 bytes) Left: The first Rhinobatos incertus (male breeding phase) tooth to be found in the lower (late Coniacian) Smoky Hill Chalk. Although not common anywhere in the Niobrara, Rhinobatos has been previously reported from the upper chalk (early Campanian) by Stewart (1990) and most recently from the middle (Santonian) by Beeson and Shimada (2004).  An unreported Rhinobatos tooth   from Kansas is in the collection of the Los Angeles County Museum of Natural History. (see Everhart, 2007)

Right: A total of six small Rhinobatos teeth were recovered from the same sample.

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shrhino3a.jpg (19195 bytes) Left: A male breeding phase Rhinobatos incertus (FHSM VP-16442) tooth that I recently (10/2005)  recovered from a matrix sample collected by Chuck Bonner from just below the Pierre Shale-Smoky Hill Chalk contact. This tooth provides evidence of a nearly continuous occurrence of Rhinobatos from the Albian Kiowa Shale through the Lower Campanian Smoky Hill Chalk.

Right: A second, slightly larger Rhinobatos incertus tooth (VP-16441)  recovered from the same sample.

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Superorder: Holocephali

Order:           Chimaeriformes

Family: Callorhynchidae    -     Eumylodus laqueatus Leidy, 1873

ratfisha.jpg (18554 bytes) LEFT: A modern ratfish (Chimaea monstrosa Linnaeus 1758 - also called the Rabbit fish). Although many species of these fish have been around since the Triassic, their remains are relatively rare in the fossil record.  Most chimaeroid remains are limited to teeth (tritors), fin spines and paired jaw plates because they are a cartilaginous fish like the sharks described above. Although the jaw plates (see below) appear to be bone, they are actually composed of calcified cartilage. Modern ratfishes typically use their crushing dentition to feed on hard shelled prey. The tritors are composed of hypermineralized cartilage. 
Ratfish1a.jpg (15317 bytes) LEFT: Recently discovered jaw elements (FHSM VP-16685) of a Late Cretaceous chimaeroid (ratfish) - Eumylodus laqueatus Leidy, 1873 (REVERSE VIEW). This is only the second known specimen of a ratfish from the Smoky Hill Chalk. A dorsal fin spine was discovered by H.T. Martin along Hackberry Creek (Gove County) about 1900, and was sold to the British Museum of Natural History (BMNH P10343, see Stahl, 1999, fig. 143). Ratfish have three pairs of jaw plates (vomerines, palatines and mandibulars).  The specimen was identified almost simultaneously by Ken Carpenter (Denver Museum of Natural History) in August, 2006, and also by David Parris (New Jersey Museum of Natural History) and David Cicimurri (Bob Campbell Geology Museum, Clemson University).

PUBLICATION: Cicimurri, D. J., D. C. Parris and M. J. Everhart. 2008. Partial dentition of a chimaeroid fish (Chondrichthyes, Holocephali) from the Upper Cretaceous Niobrara Chalk of Kansas, USA.  Journal of Vertebrate Paleontology 28(1):34–40.

Class Osteichthyes (Bony Fish):


Bony fish (including teleosts) were evolving rapidly at this time and beginning to produce the variety of modern shapes and sizes that we see today. Some of the largest bony fish that ever swam were in the Western Interior Sea during the Cretaceous Period and have left abundant remains. Many times, however, only the tail of the fish remains as evidence of the leftovers from a larger predator's meal.


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Most of the remains that are found were from relatively large fish that must have been high in the food chain. None of the smaller fish, which must have existed in great abundance to support the other fish, mosasaurs, plesiosaurs, Pteranodons and hesperornids are well represented in the fossil record either because their skeletons were too small to be preserved except under special conditions, and more likely, they were usually consumed by larger predators.


LEFT: Xiphactinus audax and squid - Copyright © Doug Henderson; used with permission of Doug Henderson


Subclass Actinopterygii

    Division Halecostomi

        Suborder Pycnodontidae


Micropycnodon kansasensis


Pycnodontids: Small fishes, similar in many respects to the modern Parrot Fish, with crushing teeth that look like small round pegs.


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LEFT: Apparently, this fish "nibbled" on the invertebrates that were attached to inoceramid shells at the bottom of the ocean, although they are more commonly found and may have preferred much shallower water near shore. Pycnodontids are found from the Early Cretaceous Kiowa Shale (Albian) through the lower half of the Smoky Hill Chalk (Coniacian).


RIGHT: A well preserved and fairly complete specimen of Micropycnodon kansasensis (KUVP 7030) in the collection of the University of Kansas.

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LEFT: A photograph of two pycnodont jaw plates. The one at far left was found by Shawn Hamm; the other was found by Pam Everhart in 1991. Two additional pictures of the right, lower jaw of the 1991-84 pycnodontid (Micropycnodon) are here: (Photo 1Photo 2)


LEFT: A left prearticular toothplate of a juvenile pycnodont collected by Keith Ewell in 2004 from Trego County. The white color of some of the teeth indicates oxidation of the enamel due to weathering. (Scale = mm)

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KU127042-1a.jpg (16674 bytes) LEFT: A nearly complete specimen of Micropycnodon kansasensis (KUVP 127042) in the collection of the University of Kansas Museum of Natural History found by Laverne "Duffer" Mauck in southeastern Gove County in 1994 near Hattin's Marker Unit 4. Head is to the left.

RIGHT: A close up of the spiky scales covering ventral side (keel) of this fish. The spines are made up of a enamel like substance called ganoin which covers the scales and dermal bones of ganoid fish (Also visible on the type and KUVP 7030) 

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Division Ginglymodi


Lepisosteus sp. (Gars)


Gar fish - A single, disarticulated specimen (KUVP 36243) was reported by Wiley and Stewart (1977) from the lower chalk (Late Coniacian, zone of Protosphyraena perniciosa) of Trego County, KS.


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LEFT: FHSM VP-5114 -These large Lepisosteus sp. scales were collected from the Kiowa Formation of Kiowa County and are Albian in age.  Scales of Lepisosteus sp. are also known from the Buildex Quarry in McPherson County, along with the teeth of Lepidotes sp.


RIGHT: Distinctive crowns of three teeth from the Lepisosteus sp. (KUVP 36243) remains described from Trego County by Wiley and Stewart (1977).

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Division ?

    Family Pachycormidae ?


Genus Protosphyraena Leidy, 1857




A large, fairly common fish that would have looked somewhat like a modern swordfish with a sword-like beak and long, sickle shaped fins. They grew to large size. The pectoral fins and shoulder bones are most frequently preserved as fossils of this genus, and in several species the fins have a distinct saw toothed, leading edge. The genus includes: Protosphyraena perniciosa (Cope, 1874); Protosphyraena nitida (Cope, 1872); Protosphyraena tenuis Loomis, 1900; Protosphyraena gladius (Cope, 1873), and Protosphyraena bentonianum (Stewart, 1898).


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LEFT: This is the most recent concept of what  Protosphyraena may have looked like, based on the first  recovery of a nearly complete specimenAlthough lacking the anterior portion of the skull, it is the most complete specimen of Protosphyraena ever found, and provides valuable information regarding the characteristics of this fish. The fish is rather short for the size of it's long, saw-toothed pectoral fins and large nearly vertical caudal fin. The pelvic fins are long and streamer-like, and emerge just behind the pectoral fins.

 RIGHT: The tail of a Protosphyraena sp. specimen (KUVP 49419) in the University of Kansas collection.  Closer view HERE

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The skull, with a sharp rostrum (nose) is also found preserved in the chalk and is readily recognized by the flat, blade-like teeth. The  anterior teeth in both jaws point forward. The remaining skeleton was made up of poorly ossified bone and has not been found preserved, except for the hypural bone of the caudal fin.


RIGHT: The sharp snout of a Protosphyraena nitida skull (from Hay, 1903). These relatively heavy pieces of bone are fairly common in the chalk.  The heavy scapulo-coracoid and pectoral fins are usually found preserved separately.


Protosphyraena perniciosa (Cope, 1874)


A large predatory fish with long pectoral fins that had a sharp, serrated leading edge. Fins and elements of the pectoral girdle are found most often. Found in lower 1/3 and extreme upper portions of the chalk.



LEFT: A complete set of Protosphyraena perniciosa fins (FHSM VP-80) that are on exhibit in the Sternberg Museum, Hays, Kansas. These were collected in the lower Smoky Hill chalk of Ellis County by George F. Sternberg. Each fin measures over 36 inches, and the complete mount measures 77 inches with the scapulo-coracoid bones. (CLICK TO ENLARGE)


The original name given to this fish by Cope in 1874 was Ichthyodectes perniciosus. He did not recognize that it was in the genus Protosphyraena, named by Leidy in 1857.

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LEFT: A complete skull of Protosphyraena sp. (FHSM VP-81) Probably P. perniciosa) that is on exhibit in the Sternberg Museum, Hays, Kansas. This specimen was collected and prepared by George F. Sternberg, and measures 15 inches from the base of the skull to the tip of the rostrum. Note the forward facing teeth in both the upper and lower jaws.


Protosphyraena nitida (Cope, 1872)


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LEFT: Originally called Erisichthe nitida by Cope in 1873, Protosphyraena nitida was a medium sized predatory fish with pectoral fins characterized by fine lines perpendicular to the leading edge of the fin. Found in lower 1/3 of chalk. Mike and Pam Everhart  collected the only known complete skull and fins of this fish. The specimen (LACM 129752) is now at the Los Angeles County Museum of Natural History.

EPC1994-04a.jpg (28870 bytes) LEFT: (EPC 1994-04) Protosphyraena nitida fins do not have the jagged, saw-toothed edges found in P. perniciosa. Instead they have fine grooves at right angles to the edge of the fin (CLICK HERE FOR CLOSE-UP).
EPC2003-14a.jpg (22837 bytes) LEFT: An associated partial skull and pectoral fin of a very small (about 3 ft or 1 m) Protosphyraena nitida (EPC 2003-14) that I collected in the lower chalk of southeastern Gove County (2003). This specimen is unusual both for the small size and the association of a rostrum with a pectoral fin. The specimen shows evidence of being scavenged by a shark, probably Squalicorax falcatus.

Protosphyraena tenuis (Loomis, 1900)


A medium sized predatory fish with sickle shaped fins characterized by a moderately serrated leading edge. The serrations are not thickened in cross section as they are in P. perniciosa. This species appears to replace P. perniciosa above the lower 1/3 of the chalk.

EPC1994-15a.jpg (22894 bytes) LEFT: A partial pectoral fin of Protosphyraena tenuis Loomis 1900 in my collection (EPC 1994-15). CLICK HERE FOR A DETAIL OF THE EDGE OF THIS FIN.
EPC1995-38a.jpg (29655 bytes) LEFT: A partial pectoral fin of Protosphyraena tenuis Loomis 1900 in my collection (EPC 1995-38). CLICK HERE FOR A DETAIL OF THE EDGE OF THIS FIN.


Bonnerichthys gladius (Cope, 1873)


FHSM VP-212a.jpg (31008 bytes) LEFT: A very large but relatively rare fish that is found in the upper 2/3 of the chalk. Pectoral fins are massive in both length and thickness. This fish was recently determined not to a Protosphyraena (swordfish). Little was known about the skull until recently, but now it appears to be a large filter feeder like Leedsichthys from the Jurassic of England. It was redescribed and renamed by Friedman et al. (2010).  Protosphyraena gigas (Stewart, 1999) is a junior synonym of P. gladius.


Protosphyraena bentonianum (Stewart, 1898)


This species is a poorly known from the Lincoln Limestone Member of the Greenhorn Limestone (Upper Cenomanian) that was described from a single specimen collected in Mitchell County by S. W. Williston in 1897. More recently Keith Ewell and I have collected Protosphyraena teeth from below the contact of the Dakota Sandstone and Graneros Shale in Russell County. 


KUVP414B.jpg (17697 bytes) LEFT: Albin Stewart (1898, p. 27) described a new species of Protosphyraena from the Lincoln Member (Upper Cenomanian) of the Greenhorn Limestone that had been found by S. W. Williston in 1893 in southern Mitchell County.  Stewart named the new species Protosphyraena bentonianum. The specimen (KUVP 414) consists of a 20 cm (8 in) rostrum and several unidentifiable bone fragments.  Although the type material does not contain teeth, it is likely that the teeth in the photo at above left are also from P. bentonianum. The inset is a close-up of the surface texture of the snout which Stewart believed to be distinctive for the species. However, the same texture also appears on Protosphyraena snouts from the Smoky Hill Chalk.

Note that the species name was first published as "bentonia" in 1898 by Stewart, but then was noted to be a typographical error by Stewart (1900).  

decprota.jpg (9869 bytes) LEFT: A recently collected (2005) specimen of Protosphyraena perniciosa(?) / P. bentonianum from the Fort Hays Limestone of Jewell County (private collection). Note the same surface texture on the snout as noted above.



Subdivision Teleostei (teleost fish)

    Order Ichthyodectiformes

        Family Ichthyodectidae




A family of medium to very large, primitive bony fish; includes Gillicus, Ichthyodectes and Xiphactinus.


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LEFT: All ichthyodectid fishes were covered with oval-shaped scales that were wider than they were long. Sometimes these scales are preserved in the chalk, unassociated with other vertebrate remains. The scale at left measures about 3.5 cm (1.5 in.) by 2 cm (the anterior part of the scale is in the lower part of the picture). From the large size of this scale, it is most likely from a Xiphactinus or an Ichthyodectes.

Xiphactinus audax:


Xiphactinus was the largest bony fish of the Late Cretaceous, reaching lengths of nearly 20 feet. These fish must have been the terror of the seas as far as anything smaller was concerned. Nicknamed the "Bulldog" fish because of the upward thrust of it's lower jaw, the fish would have been able to swallow very large prey, and this sometimes got them into trouble.

RIGHT: Xiphactinus audax and Gillicus


A number of complete specimens of Xiphactinus have been found with smaller fish (Gillicus) inside. This could indicate that death was somehow caused by the smaller fish but we will probably never know. The specimen shown above (FHSM VP-333 and VP-334) is in the Sternberg Museum at Fort Hays State University and was collected by George F. Sternberg in 1952. Click on the picture to see larger image. Xiphactinus page HERE.


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While the name Xiphactinus has been around since 1870, there has been considerable confusion over which name is actually correct: Xiphactinus audax (Leidy, 1870) or Portheus molossus (Cope, 1872).  Joseph Leidy named the fish from a fragment of a pectoral fin (USNM 52; at left) which was donated to the Smithsonian by Dr. George M. Sternberg a year before E. D. Cope gave the name of Portheus molossus to a collection of several nearly complete specimens found near Fort Wallace.  Leidy won by virtue of being first to publish, but Cope's name was more popular and is still in use in many collections of Cretaceous fish

The lower jaw of Xiphactinus audax (about 10 inches in length). The teeth of this fish are very large and are generally unequal in size.


The vertebrae of Xiphactinus audax. These are the largest bony fish vertebrae found in the Smoky Hill chalk and may reach 3 or more inches in diameter.  The vertebrae are spool-like, without ornamentation. This fish has no living relatives in today's ocean but would probably have looked much like a modern tarpon.


Pictures of two Xiphactinus audax vertebrae are HERE and HERE (scale - mm)

The pectoral fins of Xiphactinus were large and heavy, and were almost entirely made of solid bone.



Ichthyodectes ctenodon:


A larger ichthyodectid fish with heavy jaws and small, equally sized teeth (see jaws below). It grew to lengths of 8 to 10 feet and was a predator on smaller fish (see Everhart, et. al., 2010). The complete specimen (below) is a juvenile and is one of the smallest known reasonably complete examples of this species.  (BELOW) Note that the bone on top of the head is the supraoccipital and was where major muscles were attached.. it is NOT at horn or spike and

would not be visible in the living fish. 


LEFT:  USNM 12358 is one of the most complete examples of Ichthyodectes ctenodon ever discovered. Only the tail is restored. It was collected by George F. Sternberg and Myrl V. Walker in 1931 (GFS 4931). The dig was photographed and some of the photos are in the archival collection of the Forsyth Library at Fort Hays State University, Hays, Kansas. 

LEFT: FHSM VP-760 - About 49 inches long, collected by Marion Bonner in Logan County, KS - (Click here for skull).  Bardack (1965) notes that the fins and tail are restored and that  the "ventral ends of pleural ribs are not preserved, giving this fish an appearance of exaggerated length in proportion to its depth." The spike on the head (supraoccipital) was added during preparation. (File photo, Sternberg Museum of Natural History).

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Vertebrae are large, spool shaped, and without ornamentation. Ichthyodectes is very similar to Gillicus in shape, but somewhat larger.


LEFT: The skull of Ichthyodectes ctenodon in the Museum of Natural History at The University of Kansas.


The lower jaw of Ichthyodectes ctenodon showing the uniform size of the teeth.


Gillicus arcuatus


A medium-sized, very elongate predatory fish with thin, blade like jaws having only a single row of very tiny teeth. These fish reached a length of 5 or 6 feet and may have fed on plankton or other small food material filtered from the water around them, but more likely ate smaller fish. Their remains are fairly common. The genus was named by O. P. Hay (1898) in honor of Dr. Theodore Gill of the National Museum (USNM - Smithsonian).  It had originally been described in 1875 and named Ichthyodectes arcuatus by E. D. Cope.


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LEFT: A fairly complete specimen of Gillicus arcuatus - FHSM VP-420 - 1.3 m

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LEFT: A very large specimen of Gillicus arcuatus in the exhibits of the University of Kansas Museum of Natural History. One of the best known examples of this fish is found inside the body cavity of a much larger Xiphactinus audax that is on exhibit at the Sternberg Museum. (Click here for a view of a complete Gillicus in the Denver Museum of Nature and Science)


Although these fish are sometimes considered to be 'toothless'... and "even filter-feeders", a close look at their jaws will show that they are well-armed with many small teeth. A six foot long fish with a big mouth is quite capable of taking smaller fish and squid, with or without an array of impressive teeth.


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These three Gillicus skulls were prepared by Chuck Bonner and are on exhibit in the Keystone Gallery, 25 miles south of Oakley, in Logan County, Kansas.

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Subfamily Saurodontinae:


Saurocephalids (Saurodon leanus Hays, Saurocephalus lanciformis Harlan and Prosaurodon pygmaeus Loomis) - A medium sized predatory fish with a sword-like projection from the lower jaw (pre-dentary). Teeth are small, equal-sized and blade shaped.

LEFT: One on the most on complete specimens of Saurodon leanus known, this specimen was collected by Marion Bonner from the Smoky Hill Chalk of Western Kansas and sold to the Los Angeles County Museum of Natural History. This photos shows a cast of the specimen in the Rocky Mountain Dinosaur Resource Center; a closer view of the skull is here. (Photo by Anthony Maltese, used with permission)

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LEFT: Painting of Saurodon leanus by Charles Bonner, Keystone Gallery

RIGHT: The skull of Saurodon leanus, showing the unusual extension of the lower jaw (predentary).


The type specimen of Saurocephalus lanciformis Harlan 1824 is the only surviving fossil from the Lewis and Clark Expedition of 1803-4. It was found in NW Iowa and is now in the Academy of Natural Sciences of Philadelphia (ANSP 5516).


See also this historical paper: Harlan, R., 1824. On a new fossil genus of the order Enalio sauri, (of Conybeare). Journal of the Academy of Natural Sciences of Philadelphia, Series 1, 3(pt. 2):331-337; pl. 12, figs. 1-5.

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LEFT: The skull of Saurodon leanus Hays in the exhibits of the Sternberg Museum of Natural History. (FHSM VP-168)

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LEFT: The lower jaws / predentary, and partial right upper jaw / premaxilla of Saurodon leanus (V-968) from the Fryxell Geology Museum at Augustana College in Rock Island, Illinois.  The partial lower jaw is 20.7 cm in length. This specimen was collected in 1941 by George Sternberg (GFS 10841) from the Smoky Hill Chalk in western Gove County along the Smoky Hill River.  Sternberg originally sold the specimen to the University of California Museum of Paleontology where it was curated as UCMP 57555.  It was subsequently obtained by the Fryxell Museum in an exchange of specimens (Pers. comm., 2012, Susan Kornreich Wolf, Associate Curator, Fryxell Geology Museum ).  Here is a more recent photo.


As described by George F. Sternberg: "#10841 Saurodon: The left upper jaw with premaxillae in place. Both [of] the lower jaws with predentary in place. Total length 8 1/4". The inside of the upper jaws shows with both lowers in place. A few teeth are missing. Size of the slab 11" X 6". Bone and teeth good. A few teeth missing. Unprepared."

LEFT: The predentary of Saurocephalus lanciformis, adapted from Bardack and Sprinkle (1969, fig.1), based on an unnumbered specimen in the Museum of Comparative Zoology, Harvard University. Note that the predentary is much smaller and shorter in S. lanciformis than in S. leanus

RIGHT: Two S. lanciformis predentarys from the upper chalk (Early Campanian), Logan County, Kansas, in left lateral and posterior view.  


Order Tselfatiiformes

    Family Plethodidae




Bananogmius evolutus (Cope)


Pentanogmius evolutus (Cope) is one a diverse family of plethodids from the Late Cretaceous of Kansas. The remains of plethodids are a fairly common in the Smoky Hill Chalk. The family Plethodidae includes the genera: Pentanogmius (originally "Anogmius" evolutus, then Bananogmius), Niobrara, Zanclites, Luxilites, Syntegmodus, Bananogmius, Martinichthys, Thryptodus, and Plethodus. The species "Bananogmius evolutus" was recently removed from the genus Bananogmius and renamed Pentanogmius evolutus by Taverne).


 A medium sized fish (4 to 6 feet) with a narrow, but deep body (like an Angel Fish) and jaws that had small, comb-like teeth. Most of the skeleton was bone and preserved well. A commonly found fossil is the palatine bone from the roof of the mouth. These have a pebbled surface that served as the base for hundreds of small, sharp teeth.

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Pentanogmius was a medium to large sized fish (4 to 6+ feet) with a narrow, but deep body (somewhat like a modern Angel fish) and jaws that had small, comb-like teeth. Most of the skeleton was bone and preserves well. A commonly found fossil is the palatine bone from the roof of the mouth. These have a pebbled surface that served as the base for hundreds of small, sharp teeth.


E.D. Cope originally named the genus "Anogmius" in 1877, but that genus name had already been used (e.g. pre-occupied).


LEFT: DMNS 57159 - A large and nearly complete Pentanogmius evolutus (Cope) discovered by Chuck Bonner in the Smoky Hill Chalk of western Kansas. The tail is reconstructed.

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LEFT: The exhibit specimen of Bananogmius (Pentanogmius) evolutus (FHSM VP-2182) in the collection of the Sternberg Museum. Although not fully displayed in this photograph of the reconstruction, the fish had a large, sail-like dorsal fin.

LEFT: Bananogmius (Pentanogmius) evolutus  FHSM VP-2117) collected by Marion Bonner in the Sternberg Museum of Natural History. The caudal fin (tail) is reconstructed. A close up view of the skull is HERE. A view of the teeth on the lower jaw of another specimen is HERE

KUVP27815a.jpg (20632 bytes) LEFT: The skull of a large Bananogmius (Pentanogmius) evolutus - KUVP 27815 in left lateral view collected from the Pierre Shale in 1971 by the Orville Bonner party.  

RIGHT: The type specimen of Bananogmius ellisensis, preserved uncrushed in an ironstone concretion from the Blue Hill Shale Member of the Carlile Shale, Ellis County, Kansas (Middle Turonian in age).

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Complete skulls of this strange, "battering-ram" fish are found rarely in the chalk, but mostly, only the rostrum at the tip of the skull survives. Unfortunately, the type specimen from the Kansas Chalk was destroyed in Germany during World War II.


LEFT: The skull of Thryptodus was heavily built, with a rostrum (the ethmoid bone) that is very massive.  Another unusual feature is the flat surface at the anterior end. How was this fish using its head?   Click HERE for a drawing of the skull of Thryptodus zitteli (from Loomis 1900). In Kansas, these fish are only found in the lower 1/4 of the Smoky Hill Chalk and apparently become extinct by the end of the Coniacian stage (about 86 mya).




These fish are somewhat mysterious in that they are only known to occur in Kansas, and only in a relatively narrow section of the chalk. There are only two partial skulls of these fish known and few vertebrae. One of them (KUVP 497) is shown here.

kuvp497a.jpg (25626 bytes) LEFT: The type specimen of Martinichthys brevis (KUVP 497) in the collection of the Natural History Museum of The University of Kansas, and the only known, reasonably complete skull of Martinichthys. This is the only known publication of a picture of this skull since it was first described and published with photographs in 1926 by C. E. McClung (A drawing of this skull was published by Taverne, 2000).  Note that there are five vertebrae associated with the skull.  (Scale = cm)
kuvp498a.jpg (10273 bytes) LEFT: The skull of the type specimens of Martinichthys ziphioides (KUVP 498) in the collection of the Natural History Museum of The University of Kansas. There are 21 vertebrae associated with this specimen. (Scale = cm)

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LEFT: Martinichthys "noses" or "snouts" are usually the only part of the fish that is preserved. This is a drawing the type specimen (AMNH FF_2131) in the American Museum of Natural History (adapted from O.P. Hay, 1903). Note that both Cope and Hay described / figured this specimen "upside-down."


The specimen was collected by Charles H. Sternberg in June, 1877 and  described by Cope:   Cope, E. D., 1877. On the genus Erisichthe. Bull. U. S. Geol. and Geog. Surv. iii, article xx. pp. 821-823.

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These fish are usually represented in the fossil record only by a hard, blunt nose or rostrum. These rostra are often worn off at an angle as if the fish was using it to hammer at a hard substrate such as shells. The teeth are very small and comb-like in appearance. These fish only occur in a narrow zone in the lower 1/3 of the chalk. They are relatively rare (found only in Kansas) and there are less than 85 examples in collections. Most of the specimens are in the collections of the Sternberg Museum and the Museum of Natural History at the University of Kansas.


LEFT: FHSM VP-15567 - A recently discovered (2003), well-preserved Martinichthys brevis rostrum from the lower Smoky Hill Chalk of Gove County, KS.


Other Plethodids:


Relatively rare and unknown.  These specimens are in the KU Museum of Natural History:


KUVP 296a.jpg (19292 bytes) LEFT: The skull of Ferrifrons rugosus Jordan 1924 (KUVP 296), the anterior portion of a complete specimen (damaged). It was almost 75 years later that this species was determined to be a plethodid (see Arratia and Chorn. 1998. Journal of Vertebrate Paleontology, 18(2): 301-314. The original specimen is 24.5 inches long and was collected by H.T. Martin about 4 miles northeast of Gove, in Gove County, KS.

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LEFT: Niobrara encarsia Jordon 1924 - This specimen was collected by H.T. Martin in Trego County. Another view of the type specimen (KUVP 179).

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LEFT: Zanclites xenurus Jordon 1924 - This specimen was collected by H.T. Martin about 1/2 mile northeast of Gove, Gove County, KS.  Another view  of the type specimen (KUVP 52)




Aethocephalichthys hyainarhinos


Headlamp Fish: A very unusual fish that is known only from the highly fused neurocrania that have been found in Mississippi, Kansas, South Dakota and New Zealand.


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LEFT: The specimen shown at left is from the Pierre Shale.   It was found by Pam Everhart in 1995, and donated to the Sternberg Museum in 2004 (FHSM VP-15577). However, at least one specimen is known from the Smoky Hill Chalk in Gove County and is in the collection of the University of Kansas.


RIGHT: FHSM VP-16466 is a new specimen collected in the Smoky Hill Chalk of Gove County, Kansas.


For more information, see: Fielitz, C., Stewart, J.D. and Wiffen, J. 1999. Aethocephalichthys hyainarhinos gen. et sp. nov., a new and enigmatic Late Cretaceous actinopterygian from North America and New Zealand, pp. 95-106, 7 fig., in Arrantia, G. and Shultze, H-P. (eds.), Mesozoic Fishes 2 - Systematics and Fossil Record.

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Order Elopiformes

    Family Pachyrhizodontidae




Larger members of this genus have a heavy skull, with large, conical, curved teeth. The remains of the jaws often have been mistaken for marine reptiles such as mosasaurs.


Pachyrhizodus minimus


A small predatory fish sometimes found complete with well preserved with scales and internal structure. 

vp-326da.jpg (40481 bytes) LEFT: FHSM VP-326 - A very complete specimen in the Sternberg Museum of Natural History; collected by G. F. Sternberg in Trego County in 1949 and described by H.W. Miller (1957): Intestinal casts in Pachyrhizodus, an Elopid fish, from the Niobrara Formation of Kansas. Kansas Academy of Science, Transactions 60(4): 399-401.
VP-2282Aa.jpg (21907 bytes) RIGHT: FHSM VP-2282, a small Pachyrhizodus minimus found by Ken Metzler in Ellis County (1968).


Pachyrhizodus caninus


 A medium sized predator, with long heavy jaws. The premaxillary of this species is relatively easy to identify because it has one or two large fangs set inside the outer row of teeth. Click here for photos of  the skull and jaws of a very large Pachyrhizodus caninus specimen (FHSM VP-2189) in the collection of the Sternberg Museum.


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ABOVE: A reconstructed skeleton of  Pachyrhizodus caninus at the Rocky Mountain Dinosaur Resource Center.


RIGHT: The right maxilla and right dentary of Pachyrhizodus caninus.


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Pachyrhizodus leptopsis


A rare species in the lower chalk, differentiated from P. caninus by the shape of the teeth. While P. caninus teeth are round or oval in cross section, P. leptopsis teeth are distinctly fluted.



Order Elopiformes

    Suborder Elopoidei

        Family Apsopelicidae


             Apsopelix anglicus (Dixon 1850)  

LEFT: Apsopelix anglicus (less than 1.5 ft) was a small, relatively uncommon fish from the Late Cretaceous, with a wide distribution. It was first described by F. Dixon in 1850.  (adapted from Teller-Marshall and Bardack 1978, Fig. 10)


RIGHT: The exhibit specimen at the Sternberg Museum. Apsopelix has been collected mostly from the lower Smoky Hill Chalk in Kansas. Specimens are usually preserved more or less complete, and often in 3-dimensions. Another Sternberg specimen is here 

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LEFT: Apsopelix anglicus (FHSM VP-2056) - a mounted specimen in the collection of the Sternberg Museum. 


RIGHT: An Apsopelix anglicus specimen (KUVP-49402) in the University of Kansas collection.  Collected by J. D. Stewart from Gove County in 1997.


For more information, see: Teller-Marshall, S. and D. Bardak. 1978. The morphology and relationships of the Cretaceous teleost Apsopelix. Fieldiana Geology 41(1):1-35.

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Superorder Neoteleostei

Order Alepisauriformes

Cimolichthys nepaholica


Cimolichthys was a common, small to medium size predatory fish. This species has a triple row of teeth on the lower jaws and premaxillae, and no teeth on the maxillae. It also has very ornate vertebra, similar to those of Enchodus, a related genus. In addition, in some specimens, large scales or scutes are preserved along the lower portion of the body. This fish may have looked much like a modern barracuda or freshwater pike. Based on two finds of Cimolichthys with the undigested remains of their final meal still inside, it is known for certain that this fish fed on Enchodus petrosus.


KU-CIM01a.jpg (35271 bytes) LEFT: A complete specimen in the Museum of Natural History at the University of Kansas.  (Click here for another specimen from the Denver Museum of Nature and Science) Additional pictures by Kenneth Jenkins here: Skull: Complete fish;   vertebrae




Fish in the genus Enchodus were small to medium sized predators. Several species are well represented throughout the chalk and are commonly found as fossils. The characteristic that is most noticeable is the presence of large "fangs" at the front of the upper and lower  jaws.


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Above: A drawing of Enchodus petrosus showing the oversized fangs which were typical of these fish. The upper fangs are slightly larger than the lower ones. Note that when the jaws are closed, the large palatine fangs of the upper jaw are between the large teeth on the lower jaws.  This fish has also been called the "Saber-toothed Herring" because of the appearance of these unusually large, curved teeth.  It is important to note, however, that Enchodus is not related to Herrings and in fact, is a member of the same Order as modern Salmon. It is possible that the fish may used it's unique teeth to feed on soft bodied animals like squid (Tusotuethids). This genus of fish survived the end of the Age of Dinosaurs by several million years and is well represented in the fossil record around the world.

Typical examples of Enchodus remains that are found in the Smoky Hill Chalk. Top row: Two palatine bones with partial fangs; Bottom row: two lower jaws.


Enchodus petrosus:


This fish was the largest of the Enchodontids found in the Smoky Hill Chalk (Niobrara Fm) of western Kansas.


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LEFT: This is a picture of a fairly complete and unusually large set of Enchodus petrosus teeth found by the author in September, 2002. The fish would have been 4-5 feet long.  It is little wonder that they have been referred to (incorrectly) as the "Saber-toothed Herrings" of the Cretaceous. They are more closely related to modern salmon.


Enchodus gladiolus


Another small predatory fish, with elongated fangs on the lower jaws and the palatine bones of the skull. E. gladiolus was a small sized predator, with palatine teeth that reached two inches in length. Pectoral fins are long and delicate, and may have given the fish the capability to 'fly' in a manner similar to modern flying fish.


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LEFT: This specimen was found in the lower Smoky Hill Chalk by Pam Everhart and donated to the New Jersey State Museum.


RIGHT: The exhibit specimen of Enchodus gladiolus at the Sternberg Museum of Natural History. It appears to be missing the palatine tooth at the front of the upper jaw. These teeth were shed and replaced periodically as the fish grew larger and the length of the jaws increased.

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Enchodus dirus - A small, relatively unknown species of Enchodus.



Apateodus - The genus Apateodus (Teleostei: Aulopiformes) was first described from specimens in the English Chalk by Woodward in 1901. It is a medium sized bony fish that apparently had a worldwide distribution in the Late Cretaceous, but is only known from a few specimens with post-cranial material (also in Russia and India). A reasonable complete specimen (AMNH FF11560; Below) was collected by George Sternberg from Wildhorse Canyon in SW Trego County in 1949 and sold to the American Museum of Natural History.

LEFT: From Plate XIV in Woodward, 1901, showing two skulls of Apateodus.

RIGHT: From Plate XI in Woodward 1902, showing the teeth of Apateodus.

LEFT: Apateodus sp. (AMNH FF11560) - Description reads: "Unidentified fish skull and partial skeleton on plaster base 27 1/2" long by 14 1/2" wide. Locality Gove County, Kansas, about 2 1/2 miles due east of Castle Rock.  In Niobrara Cretaceous chalk.  Found about 1949 by George F. Sternberg." Photo by Matt Friedman; used with permission of Matt Friedman and the American Museum of Natural History.

RIGHT: Closer view of the skull of AMNH FF11560. Photo by Matt Friedman; used with permission of Matt Friedman and the American Museum of Natural History.

LEFT: A smaller specimen of Apateodus sp. (KGM0031) collected by Chuck and Barbara Bonner from the upper chalk, eastern Logan County, and displayed in the Keystone Gallery, south of Oakley, Kansas.  Detail of the vertebral column HERE. The vertebrae appear to be very similar to those of Enchodus or Cimolichthys. Photos by Matt Friedman; used with permission.

RIGHT: Detail of the jaws of the Bonner specimen. The way that the teeth are slanted forward in the jaws is unusual. Photo by Matt Friedman. 

LEFT: This specimen of Apateodus (CMC VP 6941) was collected by a friend of mine (Pete Bussen) from the upper chalk in western Logan County and donated to the Cincinnati Museum Center. The photo shows both lower jaws. The large bone at the upper left is the ceratohyal. The specimen was subsequently described as a new species (A. busseni) by Fielitz and Shimada (2009). Photographed in 2003. 

RIGHT: Detail of the dentary showing the characteristic blade-like teeth that are inclined anteriorly. Close-up HERE.

Fielitz, C. and Shimada, K. 2009. A new species of Apateodus (Teleostei: Aulopiformes) from the Upper Cretaceous Niobrara Chalk of western Kansas, U.S.A. Journal of Vertebrate Paleontology 28(3):650-658.

LEFT: Another portion of the specimen showing the anterior portion of the vertebral column and the back of the skull of the type specimen of Apateodus busseni (CMC VP 6941). Close up HERE.

RIGHT: The base of the caudal fin of the type specimen of Apateodus busseni (CMC VP 6941).



Family Dercetidae          


Dercidids (Stratodus apicalis) - A relatively rare fish with a narrow, eel like body and numerous small teeth. The picture at below left shows the long, slender vertebral column of what had been a fairly complete specimen (EPC 1990-14) as found in the lower Smoky Hill Chalk.

kuvp-23a.jpg (16540 bytes) LEFT: The disarticulated skull and jaws of a large Stratodus apicalis (KUVP 23) collected from Logan County by S. W. Williston.

RIGHT: Stratodus apicalis specimen as found in the Late Coniacian Smoky Hill Chalk (Gove County). Head is to the lower left in the photo. Hammer is 13 in / 33 cm.

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LEFT: Vertebrae and jaw fragments from the specimen at above right.


RIGHT: A close-up of the jaw showing the fine, comb-like teeth. (Scale = mm)

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Superorder Acanthopterygii

    Order Beryciformes

        Family Holocentridae Richardson (1846)

            Genus Kansius Hussakof 1929


Kansius sternbergi

Near the middle of the chalk, large Inoceramid shells have been found with schools of small Holocentrid fish preserved inside. Sometimes numbering in the hundreds, these fish were apparently trapped inside the shell when the clam died and the shell closed. Although several new species have been found inside clam shells, most have not yet been described. The remains of one species, Kansius sternbergi (below), have been found inside and outside of clam shells.

This unusual association was first noted by G.F. Sternberg in 1930 on the basis of a group of "twenty or more" small fish discovered inside an inoceramid clam shell in northern Trego County by M.V. Walker.  The fish averaged 3.5 inches long (his SP-225-30). Sternberg also mentioned a similar discovery from the Smoky Hill Chalk west of Webster, Kansas. These associations are most common in the early Santonian, near Marker Unit 8. 

See also: Stewart. J.D. 1984. Taxonomy, paleoecology, and stratigraphy of halecostome-inoceramid associations of the North American Upper Cretaceous epicontinental seaways. Unpubl. doctoral dissertation, University of Kansas, 201 pp.

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LEFT: Kansius sternbergi - A small fish (about 3-4 inches) found rarely in the Chalk. Drawing from Hussakof (1929). Pictures of the type specimen (FHSM VP-19) in the collection of the Sternberg Museum are HERE and HERE.


RIGHT: A more complete specimen of Kansius sternbergi (FFHM 1972-128) on an inoceramid clam shell in the collection of the Fick Fossil and History Museum, Oakley, KS.

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Superorder Paracanthopterygii

    Order Polymixiiformes

        Family Polymixiidae

            Genus Omosoma Costa (1857)           


Omosoma garretti  - Omosoma was described first from Europe and the Middle East, but is found in greater numbers in the Smoky Hill Chalk (ibid., p. 357). Other specimens have been collected in Gove County, but preservation of the skeleton, especially the skull, is usually poor due to the small size. This small (4-7 cm) fish was first described from Kansas by Bardack (1976) from specimens found in the inside of an inoceramid shell from the Garrett Ranch in Trego County

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LEFT: An unusual slab of chalk collected from Gove County in 1975 containing part of a school of Omosoma garretti FHSM VP-5115) on exhibit at the Sternberg Museum.  


RIGHT: A detail from the Platyceramus platinus shell that the VP- 5115 specimen of Omosoma garretti was laying on.

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Leptecodon rectus

lepto98a.jpg (46247 bytes) LEFT:  The type specimen of Leptecodon rectus - KUVP 35.

S. W. Williston (1899, 1900) described a new genus and species of  fish (Leptecodon rectus; KUVP 35) found inside an inoceramid (Platyceramus platinus) clam shell  by H. T. Martin in 1895.


Three complete specimens of Omosoma garretti (KUVP 599016, 59017, 59018) were also associated with the new species. (See upper left of Plate 26)  Age is Santonian. 


lepto98d.jpg (9087 bytes) LEFT:  A more recent (2004) picture of the type specimen of Leptecodon rectus - KUVP 35, in the collection of the University of Kansas Museum of Natural History. The specimen is laying on a piece of inoceramid shell.
kuvp35f.jpg (20731 bytes) LEFT: It turns out that Leptecodon was an accomplished predator... both of the known specimens included stomach contents... the remains of a smaller Omosoma garretti was found inside the above specimen. Figure adapted from Figure 5 in: Stewart, J. D., 1990. Niobrara Formation symbiotic fish in inoceramid bivalves. p. 31-41 In S. Christopher Bennett (ed.), 1990 Society of Vertebrate Paleontology Niobrara Chalk excursion guidebook. Museum of Natural History and the Kansas Geological Survey, Lawrence, KS.

See also:Stewart. J.D. 1984. Taxonomy, paleoecology, and stratigraphy of halecostome-inoceramid associations of the North American Upper Cretaceous epicontinental seaways. Unpubl. doctoral dissertation, University of Kansas, 201 pp.


Caproberyx sp.

kuvp345a.jpg (16172 bytes) LEFT:  A section of inoceramid shell (Platyceramus platinus)  with the disarticulated remains of two or three small fish (Caproberyx sp.: KUVP 345) inside. Collected by H. T. Martin from the Smoky Hill Chalk (Gove County) in 1915.


Order Anguilliformes
    Family Urenchelyidae 
Genus Urenchelys Woodward
Urenchelys abditus,
Wiley and Stewart, 1981

LEFT: This is only the second specimen of a Urenchelys abditus (a tiny, 3-inch long eel) ever collected from the Smoky Hill Chalk of western Kansas. It was discovered inside a inoceramid clam shell in southeastern Gove County in 2010 by Kris Super. It was first described as a new species by Wiley and Stewart in 1981 (CLICK TO ENLARGE):

Wiley, E.O. and J.D. Stewart. 1981. Urenchelys abditus, new species, the first undoubted eel  (Teleostei: Anguilliformes) from the Cretaceous of North America. Journal of Vertebrate Paleontology 1(1):43-47.



Subclass Saurcopterygii




Until recently, there were no coelacanths known from the chalk. Coelacanths are well represented in the English Chalk (below RIGHT). The discovery of a partial skull by Pam Everhart in 1990 was the subject of a presentation (Stewart, J.D., et al. 1991) at the annual meeting of the Society of Vertebrate Paleontology, and also at the Kansas Academy of Science annual meeting in 1995 (Everhart, et al., 1995). 

Since the original discovery, another set of fragmentary remains has been collected (J. D. Stewart, pers. comm. 1994). In addition, recent discoveries of the remains of a giant coelacanth in the Smoky Hill Chalk of Lane County, Kansas have been assigned to a genus and species (Megalocoelacanthus dobiei) first reported from Alabama by Schwimmer et al. in 1994 (see also Schwimmer 2006). The latest report by Dutel et al (2011) describes a fairly complete and well preserved skull in the collection of the American Museum of Natural History. 

Although coelacanths are a very old family of fish, they are not extinct as once thought and are still present in the Indian Ocean near Madagascar, and in the East Indies.

131958-4a.jpg (18912 bytes) LEFT: LACMNH 131958 - Jaws and palatoquadrate (See yellow highlighted area in figure at RIGHT (Mawsonia).    coelacan.jpg (38701 bytes)
LEFT: A reconstruction of Megalocoelacanthus dobiei adapted from Schwimmer et al. 1994, fig. 4. 

RIGHT: A partial skull of Megalocoelacanthus dobiei (RMDRC 05-018) from the Smoky Hill Chalk of Lane County, Kansas. Photo adapted from the Rocky Mountain Dinosaur Resource Center Blogspot. This specimen was subsequently acquired by the American Museum of Natural History and is curated as AMNH FF 20267 (see Dutel, et al. 2011).


Coelacanth References:


Dutel, H., Maisey, J., Schwimmer, D., Janvier, P. and Clément, G. 2011. Giant coelacanth Megalocoelacanthus dobiei from the Upper Cretaceous of North America and its bearings on the phylogeny of Mesozoic coelacanths. Journal of Vertebrate Paleontology 31(Suppment to 3):103.


Everhart, M. J., Everhart, P.A. and Stewart, J.D. 1995. Notes on the biostratigraphy of a small coelacanth from the Smoky Hill Chalk Member of the Niobrara Chalk (upper Cretaceous) of western Kansas. Kansas Academy of Science, Transactions 14(Abstracts):20.


Schwimmer, D. 2006. Megalocoelacanthus dobiei: Morphological, range and ecological descriptions of the youngest fossil marine coelacanth. Journal of Vertebrate Paleontology 26(Supplement to 3):122A.


Schwimmer, D.R. 2009. Giant coelacanths as the missing planktivores in southeastern Late Cretaceous coastal seas. Geological Society of America, 58th Annual Meeting, Paper No. 3-1 (abstract).


Schwimmer, D. R., Stewart, J.D. and Williams, G.D. 1994. Giant fossil coelacanths of the Late Cretaceous in the eastern United States. Geology. 22:503-506.


Stewart, J.D., Everhart, P.A. and Everhart, M.J. 1991. Small coelacanths from Upper Cretaceous rocks of Kansas. Journal. Vertebrate Paleontology 11(Abstract, suppl. to 3):56A.



Rare fish - There are several genera and species of fossil fish that are sometimes represented only by few remains or even just a single specimen from the chalk. These include:  Trachichthyoides, Luxilites striolatus (KUVP 295), Paraliodesmus, and Belonostomus.


References on Sharks and Fish in the Western Interior Sea


Continued on next page.................................     A Field Guide to Fossils of the Smoky Hill Chalk: Part 3 - Marine Reptiles


Credits - Earl Manning and many others have provided me with information regarding the classification of fishes from the Smoky Hill Chalk