Lucas, S. G. and Sullivan, R.M., eds., 2006, Late Cretaceous vertebrates from the Western Interior.

New Mexico Museum of Natural History and Science Bulletin 35.

185
CAUDAL FIN SKELETON OF THE LATE CRETACEOUS LAMNIFORM SHARK,
CRETOXYRHINA MANTELLI, FROM THE NIOBRARA CHALK OF KANSAS

KENSHU SHIMADA1, STEPHEN L. CUMBAA2, AND DEANNE VAN ROOYEN3

1
Environmental Science Program and Department of Biological Sciences, DePaul University, 2325 North Clifton Avenue, Chicago, Illinois 60614; and Sternberg
Museum of Natural History, Fort Hays State University, 3000 Sternberg Drive, Hays, Kansas 67601;
2
Paleobiology, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4, Canada;
3
Department of Earth Sciences, Carleton University, 2240 Herzberg Laboratories, 1125 Colonel By Drive, Ottawa, Ontario K1S 5B6, Canada.

AbstractThe caudal fin morphology of the Late Cretaceous lamniform shark, Cretoxyrhina mantelli (Agassiz),
was previously inferred from scale morphology, which suggested that it was capable of fast swimming. A specimen
from the Niobrara Chalk of western Kansas is described here and offers new insights into the morphology of the
caudal fin of the taxon. The specimen preserves the posterior half of the vertebral column and a series of hypochordal
rays. These skeletal elements exhibit features suggesting that C. mantelli had a lunate tail and a caudal peduncle with
a lateral fluke. The specimen also supports the idea that the body form of C. mantelli resembled that of the extant
white shark, Carcharodon carcharias (Linneaus). Given a total vertebral count in Cretoxyrhina mantelli of about
230, this specimen suggests that the transition between precaudal and caudal vertebrae was somewhere between the
140th and 160th vertebrae. The estimated total body length of the specimen described here ranges from 640 cm to 700
cm, marking the largest C. mantelli individual estimated to date. New skeletal data from the specimen further supports
the view that C. mantelli was an active shark capable of fast swimming.

INTRODUCTION
Cretoxyrhina mantelli (Agassiz) was a Late Cretaceous lamniform
shark that lived in CenomanianCampanian seas worldwide, including the
Western Interior Seaway of North America (e.g., Cappetta, 1987; Siverson,
1992, 1996; Shimada, 1997d). The species is represented chiefly by its
teeth, but some reasonably complete skeletons of the species are known
from the late Coniacian Santonian portion of the Smoky Hill Chalk Mem-
ber of the Niobrara Chalk in western Kansas (Shimada, 1997b). Those
skeletal remains suggest that large individuals of C. mantelli measured about
5 to 6 m in total length and possibly had a body form similar to the modern
great shark, Carcharodon carcharias (Linnaeus) (Shimada, 1997b). The
fossil record demonstrates that Cretoxyrhina mantelli fed on large marine
vertebrates (e.g., teleosts, sea turtles, mosasaurs, and plesiosaurs: Shimada,
1997c; Shimada and Everhart, 2004; Shimada and Hooks, 2004; Everhart,
2004, 2005a) and possibly scavenged bloat-and-float carcasses of fully
terrestrial vertebrates (e.g., nodosaurid dinosaurs: Everhart and Hamm,
2005).
Despite the wealth of information about the paleobiology of
Cretoxyrhina mantelli (e.g., Shimada, 1997a, 1997b, 1997c), there are
still many unresolved questions. The morphology of its caudal fin is one
such gap in our knowledge. The previously suggested morphology, and the
estimated starting point of the caudal fin in C. mantelli (Shimada, 1997b),
were based on a series of assumptions. Shimada (1997b) demonstrated
that C. mantelli had keeled placoid scales with an average interkeel dis-
tance of approximately 45 microns. Because this interkeel distance is com-
parable to that in scales of some extant fast-swimming sharks, Shimada
(1997b) inferred that C. mantelli was also capable of fast swimming. This
evidence, combined with the fact that C. mantelli had a conical head, led
Shimada (1997b) to consider that the species had a stout fusiform body
with a lunate caudal fin for efficient hydrodynamic propulsion. Given that
the fossil species had a total vertebral count of approximately 230, Shimada
(1997b) suggested that the caudal fin of C. mantelli could have begun at
about 133rd vertebra, on the basis of comparisons with some extant fast-
swimming lamniform sharks with lunate tail fins (e.g., lamnids: Lamna
Cuvier, Isurus Rafinesque, and Carcharodon Smith).
The Canadian Museum of Nature (CMN) in Ottawa, Ontario, houses
a putative Cretoxyrhina mantelli specimen, CMN 40906, which consists
of a vertebral column and some additional skeletal elements (Fig. 1). CMN
40906 is noteworthy because of its large size and features that provide new
insights into the shape of the sharks tail. The purpose of this paper is 1) to
describe the morphology of the specimen, and 2) to discuss the caudal fin
morphology of C. mantelli and its paleoecological significance. For com-
parative purposes, fossil specimens in the following institutions are referred
to in this paper: Sternberg Museum of Natural History, Fort Hays State
University (FHSM), Hays, Kansas; and the vertebrate paleontology collec-
tion of the University of Kansas Museum of Natural History (KUVP),
Lawrence.
SYSTEMATIC PALEONTOLOGY
Class Chondrichthyes
Subclass Elasmobranchii
Order Lamniformes Berg, 1958
Family Cretoxyrhinidae Glikman, 1958
Genus Cretoxyrhina Glikman, 1958
Cretoxyrhina mantelli (Agassiz, 1843)
MaterialCMN 40906 (Figs. 15), a string of caudal and poste-
rior precaudal vertebrae with hypochordal rays and placoid scales.
Horizon and localityThe specimen was collected by G. F.
Sternberg from the Niobrara Chalk (Upper Cretaceous: see Hattin, 1982)
in western Kansas. It was sold to the Geological Survey of Canada, from
which the museum originated, in 1912.
Because exact biostratigraphic data were not available to constrain
the age of the specimen within the Niobrara Chalk, chalk samples were
taken from the matrix for stratigraphically diagnostic foraminiferans. Our
result shows that four taxa dominate the assemblage. Heterohelix globulosa
(Ehrenberg), with a Campanian Maastrichtian range, is the most abun-
dant. The other key taxa are: Archeoglobigerina cretacea (dOrbigny),
with a range from middle Coniacianearly Maastrichtian; Rugoglobigerina
rugosa (Plummer), known from the early Campanian; and Whiteinella
centennialensis Frerichs, which occurs in late Santonianearly Campanian
strata (Pessagno, 1967; Frerichs et al., 1975; Frerichs, 1979). The pres-
ence of R. rugosa, which makes up about 10% of the individuals in our
sample, strongly suggests that the Cretoxyrhina specimen came to rest on
the chalky ocean bottom in the early Campanian. The shark specimen is
thus from the upper part of the Smoky Hill Chalk. If so, whereas C. mantelli
is known from early Campanian deposits (e.g., Siverson, 1992), CMN
186

FIGURE 1. Photograph of CMN 40906, string of vertebrae with hypochordal rays of Cretoxyrhina mantelli (Agassiz) (anterior to the left; scale bar = 30 cm).

FIGURE 2. Line drawing of CMN 40906 (cf. Fig. 1), string of vertebrae (v) with hypochordal rays (hcr) of Cretoxyrhina mantelli (Agassiz) (anterior to the left; scale
bar = 30 cm).

40906 represents the youngest Cretoxyrhina specimen documented thus

FIGURE 3. Close-up view of vertebral centra (v23-v25: see Fig. 2) in CMN
40906 (scale bar = 2 cm).
far from the Smoky Hill Chalk of Kansas (see Stewart, 1990; Everhart,
2005b, table 13.1).
It should be noted that museum records indicate that the specimen
was found in Gove County, Kansas, but strata from the lower Campanian
(the uppermost Niobrara Chalk) are not known to occur there. Other fossil
specimens purchased by the museum from the Sternberg family at the same
time list G. F. Sternberg as the collector, and Logan County as the source.
Because outcrops with lower Campanian strata do occur just west of Gove
County in Logan County (Hattin, 1982), perhaps the collection data are
incorrect.
DescriptionA total of 109 vertebral centra are physically preserved
in the specimen (Fig. 1). The anteriormost ninety-five of these are clearly
in anatomical sequence and are in a good state of preservation. We refer to
these as v1 through v95, counting sequentially from the anteriormost
centrum in the specimen (Fig. 2). The v-numbers are placed in quotations
to highlight the fact that they are artificial assignments. There are impres-
sions of four vertebrae near v95 which are otherwise missing. Fourteen
smaller vertebrae, generally in poor condition, are on the slab, but are largely
disarticulated, and have not been assigned v-numbers. If the four missing
vertebrae and the 14 smaller vertebrae are counted, CMN 40906 consists
 187

FIGURE 4. Close-up view of putative caudal fin base, showing connection between vertebral column and hypochordal rays in CMN 40906 (scale bar = 10 cm; cf. Figs. 1,
2).

FIGURE 5. Placoid scales (four examples) of Cretoxyrhina mantelli (Agassiz) from CMN 40906 (scale bar = 50 micronsm). Sample locations are relative to vertebral
positions in Figure 2. A, Scale from dorsal to v24: top, apical view (anterior to the top); bottom, anterior view. B, Scale from above, or ventral to v66: left, oblique view
(anterior to the top left); right ,anterior view. C, Scale location same as A: lateral view (anterior to the left). D, Scale location same as B: apical view (anterior to the right).
188
FIGURE 6. Example of articulated vertebrae of Cretoxyrhina mantelli (Agassiz) (anterior to the left; scale bar = 10 cm; v 73v82 in FHSM VP-2187: see Shimada,
1997b). A, vertebrae in right(?) lateral view (note that this surface was against ocean floor when the shark skeleton was buried); B, vertebrae in dorsal(?) view (note post-
burial distortion that resulted in lateral flattening of vertebrae).
of a total of 113 vertebrae.
The centra (Fig. 3) are amphicoelous and asterospondylic with nu-
merous concentric lamellae around the primary double-cone calcification
(see Welton and Farish, 1993). The centra are well calcified, and the double-
cones are thick and are supported by many radial cartilage plates. A pair of
circular pits for the basidorsal and basiventral cartilages is observed on op-
posite sides of the periphery of each centrum. The pair with a narrower
inter-pit distance in each centrum is assumed to be the side of basidorsal
cartilages (see Shimada, 1997b), and thus representing the dorsal side of
the shark.
Most vertebrae are articulated, and the vertebral column shows a
dorsal bend at about v42 at an angle of approximately 45o (Fig. 2). The
anteriormost centra in the specimen are circular and measure approximately
88 mm in diameter and 18 mm in anteroposterior length (note that v1
and v2 are heavily damaged: Appendix 1). Centra towards the posterior
end of the specimen are slightly elongated dorsoventrally. Many vertebrae
suffer post-burial distortion (see below for further discussion), but they
generally decrease in their diameter gradually at a rate of 06 mm per five
vertebrae from v3 to v35 and from v40 to v95, except between
v35 and v40 with a decreasing rate of 15 mm. The last articulated
vertebral centrum has a diameter of approximately 19 mm. Some verte-
brae may be missing from the tip of the tail. The exact number of vertebrae
missing from the caudal tip is unknown, but it cannot be more than a very
few as the smallest of the disarticulated vertebrae has a centrum diameter
of about 6 mm.
A tabular piece of calcified cartilage extends ventrally from nearly
each vertebral centrum from v42 to v65 (i.e., a total of 2324 pieces:
Figs. 2, 4). These are interpreted to be hypochordal rays (see Compagno,
1999). The anteriormost ray measures 29 mm in length and 10 mm in
width. The length gradually increases posteriorly at about v51, where
the piece measures 150 mm in length and 15 mm in width (although the
widest is 25 mm at v49), and then gradually decreases towards the tip of
the tail.
A few patches of articulated placoid scales and many disarticulated
scales were found in the chalk matrix surrounding the specimen (Fig. 5).
The scales are of two types. The most common type (Figs. 5A, 5C) mea-
sures approximately 150200 microns in maximum dimension, and con-
sists of a large, keeled and fluted crown and a somewhat smaller root. The
crown has an oval apical surface and generally shows six to eight parallel,
U-shaped grooves oriented longitudinally, which emerge from the rounded
anterior margin. The posterior crown margin is thin and overlaps with pos-
teriorly located scales. The grooves are bounded on each side by a keel,
with an average interkeel distance of 2530 microns. The junction be-
tween the crown and root is constricted. The root base is flat and diamond-
shaped with one foramen at its center.
The second type was represented by only two scales in our samples
(Figs. 5B, 5D). These scales are more massive, with a prominent, rounded,
bulbous root perforated around the top by several foramina. Root to crown
length exceeds 400 microns. The crown is relatively flat, and sub-circular
to rounded quadrilateral in shape, with a greatest dimension of approxi-
mately 325 microns in the larger specimen. The crown ornamentation con-
sists of sinuous grooves and ridges in one specimen; the other, more worn
on top, appears more similar to the scales described above. The crown
edges appear somewhat worn and rounded in both specimens.
Taphonomic and taxonomic remarksA number of vertebrate
specimens from the Smoky Hill Chalk of Kansas in various museum col-
lections consist of multiple individuals that were artificially assembled for
aesthetic value (e.g., Bardack, 1965). In CMN 40906 described here, some
vertebrae are disarticulated and slightly displaced. However, we are confi-
dent that all skeletal components in CMN 40906 belong to one shark indi-
vidual, because 1) each pair of adjacent vertebrae are either articulated or
nearly identical in their vertebral diameter, and 2) the entire vertebral col-
umn along with the hypochordal rays is partially to completely embedded
in the original chalk matrix. The entire ventral surface of the specimen is

FIGURE 7. Examples of caudal fin (outline and skeleton) in selected extant sharks
(anterior to the left; not to scale). A, Isurus Rafinesque (Lamniformes: after Garman,
1913, pl. 63, fig. 5); B, Mustelus Link (Carcharhiniformes: after Mivart, 1879, pl.
LXXIV, fig. 6); C, Squalus Linnaeus (Squaliformes: after Mivart, 1879, pl. LXXVII,
fig. 3).
189
embedded in original matrix, as are v31-35 and v51-97. There is plas-
ter fill along the dorsal edge of v1-30 and v36-50, but as these verte-
brae remain anchored in chalk (except for two or three between v20 and
v25 which are loose, but have impressions in the chalk), there is no indi-
cation that the fill involved any artificial movement or substitution of verte-
brae by the preparators of the specimen.
Whereas the vertebral diameter generally decreases from the anteri-
orly located vertebrae towards the posteriorly located vertebrae, we note
that certain sections of the vertebral column, such as v4v8, v46
v62, and v70v83, show some irregularities in the maximum verte-
bral diameters (note that such measurements were excluded: see Appendix
1). The irregularities are not due to artificial causes (e.g., artificial arrange-
ment of vertebrae) but are due to post-burial distortion of vertebrae through
fossilization. Figure 6 shows a typical example of taphonomically flattened
articulated Cretoxyrhina vertebrae from the Smoky Hill Chalk of Kansas.
The effect of flattening of vertebrae is that the vertebral diameter of one
axis tends to extend, whereas the vertebral diameter of the other axis per-
pendicular to it tends to shorten (e.g., see middle section of vertebral string
in Fig. 6). Therefore, except for some slightly displaced vertebrae, the ar-
rangement of vertebrae in CMN 40906 is determined to reflect the original
sequence in the shark individual.
The taxonomy of extinct sharks generally depends on their dental
morphology. Thus, the identification of fossil shark specimens not found
in association with teeth is generally difficult. Nevertheless, the calcifica-
tion pattern characterized by many radial cartilage plates (i.e., platy pri-
mary exochordal radii: Compagno, 1990) is a strong indication that CMN
40906 belongs to the order Lamniformes. From the Smoky Hill Chalk of
Kansas, the following lamniform taxa are known to date: Scapanorhynchus
raphiodon (Agassiz), Johnlongia sp., Pseudocorax laevis (Leriche),
Squalicorax falcatus (Agassiz), S. kaupi (Agassiz), S. pristodontus
(Agassiz), S. volgensis (Glikman in Glikman and Shvazhaite), Cretalamna
appendiculata (Agassiz), and Cretoxyrhina mantelli (e.g., Hamm and
Shimada, 2002; Hamm et al., 2003; Beeson and Shimada, 2004; Shimada
et al., 2004; Shimada, 2005b, Shimada and Cicimurri, 2005). Where col-
lecting bias towards large vertebrates appears to exist in the fossil record of
the Niobrara vertebrates (Hamm and Shimada, 2002), the largest lamniform
shark known from the Smoky Hill Chalk is C. mantelli (see Hamm et al.,
2003). Based on comparisons with other C. mantelli specimens from the
Smoky Hill Chalk of Kansas (e.g., FHSM VP-323, FHSM VP-2184, FHSM
VP-2187, KUVP 247, and KUVP 69102), we note that the morphology
(e.g., the thickness of double-cone calcification and the organization of
radial cartilage plates) and general size of the vertebrae in CMN 40906
described here (e.g., Figs. 1, 3, 4) are identical to centra in those C. mantelli
specimens that are associated with teeth (e.g., Fig. 6). Therefore, we are
confident that, although CMN 40906 lacks teeth, it is a specimen of
Cretoxyrhina mantelli.
Shimada (1997b) noted that the average interkeel distance on the
crown of scales taken at the caudal region in another Cretoxyrhina mantelli
specimen (FHSM VP-2187) from the Smoky Hill Chalk of Kansas was
50.5 microns (note that, at other parts in the specimen, the smallest mean
interkeel distance recorded was 30 microns: Shimada, 1994, table 9). This
value is much higher than the average interkeel distance in CMN 40906
(i.e., 2530 m), but we note that the scale morphology in CMN 40906 is
consistent overall with Type A scales in FHSM VP-2187 (see Shimada,
1997b). Where sharks generally show large intraindividual variation in scale
morphology (e.g., Welton and Farish, 1993, fig. 20), the discrepancy in
interkeel distance values may be due to the fact that the sample size of
scales taken from FHSM VP-2187 (particularly at the caudal region) and
CMN 40906 was too small to see the possible overlap in interval ranges.
DISCUSSION
The outline of the caudal fin is not preserved in CMN 40906. How-
ever, the organization of preserved hypochordal rays with respect to the
preserved vertebral column (Fig. 4) offers insights into the tail morphology
of Cretoxyrhina mantelli. CMN 40906 shows that the anteriormost and
190
FIGURE 8. Tentative skeletal reconstruction of 6-m-long Cretoxyrhina mantelli (Agassiz) based on available fossil record (Shimada, 1997b; this study). Solid lines =
known parts; broken lines = inferential parts.
preserved posteriormost hypochordal rays are short, whereas the rays be-
tween them are markedly elongate. In addition, the base of the anteriormost
hypochordal rays in the specimen occurs immediately anterior to the sharp
bend (at an angle of ca. 45) that is observed in the vertebral column. If one
considers the sharp bend at face value as a dorsally directed bend of the
vertebral column that constitutes the axis of a tail, this skeletal organization
is remarkably similar to that of extant sharks with a high aspect ratio: i.e., a
lunate (symmetrical) tail with large dorsal and ventral lobes (Fig. 7A).
Thomson and Simanek (1977, table 3) listed Rhincodon Smith
(Orectolobiformes), Cetorhinus Blainville, Lamna, Isurus, and
Carcharodon (Lamniformes) as best representatives of this type of shark,
with the caudal portion of the vertebral column bent at an angle of ca.
=30. The caudal fin of other extant sharks is strongly heterocercal: i.e., an
asymmetrical tail with a much smaller ventral lobe compared to the dorsal
one (Thomson and Simanek, 1977). In these sharks, the vertebral column
shows a weak bend or virtually no bend, and the length of the hypochordal
rays is more or less uniform across the tail (Figs. 7B, 7C). Therefore, CMN
40906 supports the hypothesis that Cretoxyrhina mantelli had a lunate tail
(Fig. 8). It is noteworthy that, because Rhincodon is not a lamniform (Shirai,
1996), and because C. mantelli does not share an immediate common an-
cestry with Cetorhinus, Lamna, Isurus, or Carcharodon (Shimada, 2005a),
the similarity in caudal fin morphology between Cretoxyrhina mantelli
and these extant sharks is interpreted to be a homoplasy due to convergent
evolution. However, we note that the caudal fin of C. mantelli possibly had
a wider angle between the upper and lower lobes and a slightly shorter
lower lobe (Fig. 8) compared to the caudal fin of those extant sharks (e.g.,
Fig. 7A), because the hypochordal rays in CMN 40906 are largely directed
ventrally (Figs. 2, 4; as opposed to ventroposteriorly in those extant sharks).
We also note that, because the posteriorly located hypochordal rays appear
to be thinner and shorter as compared to those in Rhincodon, Cetorhinus,
Lamna, Isurus, or Carcharodon, the lobes of the lunate tail in Cretoxyrhina
mantelli could have been slightly narrower than the lobes in those extant
sharks.
Thomson and Simanek (1977) found that extant sharks with a het-
erocercal ( vertebral bent) angle of ca. 30, not only have a well- If one assumes that the vertebral diameter in CMN 40906 has the same
developed ventral hypochordal lobe, but also a deep fusiform body with a
conical head and a caudal peduncle bearing the lateral fluke. Thomson and
Simanek (1977) characterized such sharks as fast swimming pelagic
sharks, typified by extant lamnids, such as the mako and white sharks.
The vertebral bend in CMN 40906 is about 45, which approximates the
maximum possible angle observed in extant sharks (see Thomson, 1976).
Therefore, CMN 40906 further strengthens the idea (Shimada, 1997b)
that the body form of Cretoxyrhina mantelli (Fig. 8) resembled that of
extant lamnids, including the white shark, Carcharodon carcharias. We
assume that Cretoxyrhina mantelli also had a caudal peduncle with a lat-
eral fluke.
Quantitative studies of vertebral dimensions in modern sharks are
rare. However, inspection of published illustrations of caudal fins of extant
fast swimming lamniform sharks with lunate tails (e.g., Lamna in Mivart,
1879, pl. LXXV, fig. 1; Isurus in Garman, 1913, pl. 63, fig. 5: e.g., Fig.
7A) shows that a marked reduction in vertebral size is present at the
precaudal-caudal transition. There are two specimens of Cretoxyrhina
mantelli that preserve a series of vertebrae extending near the terminal
end: FHSM VP-2187 and KUVP 69102. A re-examination of vertebral
diameters in the posterior half of the vertebral column in these specimens
(i.e., post-v110: Appendix 2) reveals that a marked decrease in the diam-
eter between v140 and v145 in FHSM VP-2187 (i.e., 6 mm drop in that
interval as opposed to 04 mm per drop every six vertebrae) and between
v155 and v160 in KUVP 69102 (i.e., 8 mm drop in that interval as op-
posed to 04 mm drop per every six vertebrae where distortion is mini-
mal). Therefore, we here suggest that the caudal fin of C. mantelli possibly
occurred at an approximate range of v140v160.
If one assumes that the range of v155v160, which would give a
conservative caudal fin size, is equivalent to that of v35v40 in CMN
40906 (where a marked decrease in vertebral size is observed: see above),
the total vertebral count of the shark individual (CMN 40906) was at least
216. The number, 216, corresponds remarkably well to the estimated total
vertebral count of Cretoxyrhina mantelli, 230 (see Shimada, 1997b), by
considering the probable number of missing terminal vertebrae (i.e., 10
25 vertebrae: see above).
The largest measurable vertebra in CMN 40906 is v3, which has
a diameter of 88 mm. If v35 is assumed to be v140 in life, v3 would
then represent v108. In FHSM VP-2187, which is conservatively estimated
to be 500 cm TL, the diameter of v108 is 69 mm (Shimada, 1994, table 1).
size relation to the TL as the relationships between the diameter and TL in
FHSM VP-2187, CMN 40906 is 128% of FHSM VP-2187. This suggests
that the shark individual represented by CMN 40906 was possibly 640 cm
TL. Whereas this assumption of v35 representing v140 gives a conser-
vative TL estimate for CMN 40906, one may assume v35 to be v155 for

a less conservative estimate, and in this case, v3 would represent v123 in
life. The diameter of v123 in FHSM VP-2187 is 63 mm (Shimada, 1994,
table 1). Therefore, CMN 40906 is 140% of FHSM VP-2187, suggesting
that the shark could have measured 700 cm TL. The range of 640700 cm
TL for CMN 40906 represents the largest Cretoxyrhina mantelli individual
estimated to date (cf. Shimada, 1997b).
The caudal fin is the primary locomotor structure in sharks. Al-
though functional analysis of caudal fins is far from complete even for
living sharks (for review, see Lauder, 2000; Lingham-Soliar, 2005), ex-
amination of caudal fin morphology in extinct sharks is important because
it gives insights into their swimming capability, which in turn, as noted by
Motani (2002), has implications to behavior, physiology, and other aspects
of their biology. The exact outline of the body and tail of Cretoxyrhina
mantelli (Fig. 8) remains inferential at the present time. However, it is
noteworthy that the notion of C. mantelli as a fast-swimming shark,
supported here, agrees well with the inferred feeding behavior of the taxon
based on taphonomic evidence. In particular, a mosasaur vertebra speci-
men from Kansas shows signs of bone healing over an embedded
Cretoxyrhina tooth, suggesting the post-bite survival of the mosasaur indi-
vidual (Shimada, 1997c). Biting large, presumably active vertebrates, like
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ACKNOWLEDGMENTS
We thank M. Feuerstack and C. Kennedy (CMN) for assistance with
the specimen, and co-op students H. Bernatchez and M. Brire from lcole
secondaire de Casselman for their help in locating isolated denticles and
scale patches for study. A. Murray (CMN) took the SEM and overall speci-
men photos. We thank M. J. Everhart (FHSM) for his discussion on the
Niobrara stratigraphy. The senior author thanks the following present and
past FHSM and KUVP associates for allowing us access to comparative
Cretoxyrhina specimens in their care: J. Chorn, C. Fielitz, D. Miao, H.-P.
Schultze (KUVP), and R. J. Zakrzewski (FHSM). Comments made on
earlier drafts by D. J. Ward (Kent, United Kingdom) and M. D. Gottfried
(Michigan State University) as well as reviews by J. I. Kirkland (Utah Geo-
logical Survey) and D. R. Schwimmer (Columbus State University) greatly
improved the quality of this manuscript.
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APPENDIX 2.Approximate diameter (in millimeters) of vertebral centra in two nearly
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1994, table 1). Value in parentheses = estimated value from adjacent centra; value
in brackets = unreliable measurements due to severe distortion of centra.