A high-latitude euselachian assemblage from the early Turonian of Alberta, Canada

Abstract

Numerous isolated euselachian teeth were recovered from the early Turonian Kaskapau Formation situated in northwestern Alberta, Canada. This high palaeolatitude assemblage was collected from a sandstone lens along the bank of the Smoky River, and includes 16 species belonging to at least three orders, at least 11 families, and 15 genera. Here we describe Odontaspis watinensis sp. nov. and report the first Canadian occurrence of Polyacrodus sp., Scapanorhynchus sp., and Carcharias aff. C. striatula. The scarcity of benthic taxa in this assemblage supports the previous notion that bottom waters in this region of the Western Interior Seaway experienced enduring anoxic episodes. By comparing the faunal composition of this assemblage with that of middle Cenomanian Canadian assemblages, we show that seven species have a biostratigraphical range that extended across the Cenomanian–Turonian boundary in the northern region of the seaway. Of the taxa described herein, Archaeolamna ex. gr. kopingensis, Cardabiodon aff. C. ricki, Carcharias aff. C. striatula, Odontaspis watinensis, and Johnlongia parvidens have not been reported from deposits of the southernmost region of the seaway and may have been restricted to cooler waters.http://zoobank.org/urn:lsid:zoobank.org:pub:FD671818-2769-484C-B424-F5EF5490A6A9

Full text

Journal of Systematic Palaeontology, 2013
Vol. 11, Issue 5, 555–587, http://dx.doi.org/10.1080/14772019.2012.707990
A high latitude euselachian assemblage from the early
Turonian of Alberta, Canada
Todd D. Cooka∗,MarkV.H.Wilson
a, Alison M. Murraya,A.GuyPlint
b, Michael G. Newbreycand Michael J. Everhartd
aDepartment of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9, Canada; bDepartment of Earth Sciences,
University of Western Ontario, London, Ontario, N6A 5B7, Canada; cRoyal Tyrrell Museum of Palaeontology, Drumheller, Alberta
T0J 0Y0, Canada; dSternberg Museum of Natural History, Fort Hays State University, Hays, Kansas 67601, USA
(Received 22 June 2011; accepted 29 November 2011; first published online 20 December 2012)
Numerous isolated euselachian teeth were recovered from the early Turonian Kaskapau Formation situated in northwestern
Alberta, Canada. This high palaeolatitude assemblage was collected from a sandstone lens along the bank of the Smoky
River, and includes 16 species belonging to at least three orders, at least 11 families, and 15 genera. Here we describe
Odontaspis watinensis sp. nov. and report the first Canadian occurrence of Polyacrodus sp., Scapanorhynchus sp., and
Carcharias aff. C.striatula. The scarcity of benthic taxa in this assemblage supports the previous notion that bottom waters
in this region of the Western Interior Seaway experienced enduring anoxic episodes. By comparing the faunal composition of
this assemblage with that of middle Cenomanian Canadian assemblages, we show that seven species have a biostratigraphical
range that extended across the Cenomanian–Turonian boundary in the northern region of the seaway. Of the taxa described
herein, Archaeolamna ex. gr. kopingensis,Cardabiodon aff. C. ricki,Carcharias aff. C. striatula,Odontaspis watinensis,
and Johnlongia parvidens have not been reported from deposits of the southernmost region of the seaway and may have been
restricted to cooler waters.
http://zoobank.org/urn:lsid:zoobank.org:pub:FD671818-2769-484C-B424-F5EF5490A6A9
Keywords: Euselachian; Turonian; Western Interior Seaway; palaeobiogeography
Introduction
Only recently has there been a concerted effort to examine
the diversity of Late Cretaceous euselachian assemblages
from the northern region of the Western Interior Seaway
(WIS; see Case et al. 1990; Cumbaa & Tokaryk 1999;
Cumbaa et al. 2006, 2010; Underwood & Cumbaa 2010).
Cook et al. (2008) described a middle Cenomanian
assemblage from the Dunvegan Formation of northwestern
Alberta, Canada, reporting nine euselachian species from
three orders. The study contained the first Canadian records
of Johnlongia parvidens (Cappetta, 1973), Protolamna
carteri Cappetta & Case, 1999 and the ray Pseudohypolo-
phus mcnultyi (Thurmond, 1971). Prior to the publication
of this assemblage, the recovery of fossil euselachians from
Alberta focused on more southern and younger deposits
(Beavan & Russell 1999; Brinkman et al. 2004).
The Watino fossil localities discussed herein are signif-
icantly more productive than the aforementioned Dunve-
gan fossil localities, both with regards to the number
of teeth recovered and overall euselachian diversity. The
existence of this assemblage has been previously docu-
mented. Fox (1984) described a left humerus of Ichthy-
ornis Marsh, 1872 from Watino and also compiled a
∗Corresponding author. Email: tdcook@ualberta.ca
preliminary faunal list that included cf. Hybodus sp.,
cf. Ptychodus sp., cf. Squalicorax sp., cf. Cretolamna
sp., cf. Odontaspis sp. and Pristidae, gen. et sp. indet.
Later, Wilson & Chalifa (1989) described an assem-
blage of disarticulated and fragmentary bones and scales
of actinopterygians that included Belonostomus Agas-
siz, 1834, Ichthyodectes Cope, 1870, Apateodus Wood-
ward, 1901, Cimolichthys Leidy, 1857a, Dercetidae Wood-
ward, 1901 and Enchodus Agassiz, 1835. Also reported
at that time were the remains of Hybodontidae Owen,
1845, Ptychodontidae Jaekel, 1898, Rajiformes Berg, 1940
and Lamniformes Berg, 1958. Recently, a single tooth
belonging to the lamniform Cardabiodon ricki Siverson,
1999 was reported from Watino (Cook et al. 2010). This
tooth documented the first North American occurrence of
this large shark and provided evidence that Cardabiodon
Siverson, 1999 had an antitropical distribution.
Other important northern WIS euselachian assem-
blages have been reported from east-central Saskatchewan,
Canada. Case et al. (1990) described eight species from
the lower Turonian Keld Member of the Favel Formation
(see Cumbaa & Tokaryk 1999; Cumbaa et al. 2006, 2010
regarding the revised stratigraphy and age of this assem-
blage). Cumbaa et al. (2006) reported 19 species from the
C
2012 Natural History Museum
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556 T. D. Cook et al.
Carrot River and Bainbridge bone beds of the Belle Fourche
Member of the Ashville Formation. Recently, Underwood
& Cumbaa (2010) re-examined the latter locality and
reported one species of hybodont shark, three species
of Ptychodus Agassiz, 1835, ten species of neoselachian
sharks and two species of batoids.
Although these localities have provided us with some
knowledge of the diversity of euselachian taxa present
in the northern parts of the WIS during the Late Creta-
ceous, there is still considerable work to be done. Here
we provide a detailed examination of an early Turonian
euselachian assemblage from the Canadian region of the
seaway (palaeolatitude 59◦N; PLATES Program 2010) and
compare its composition with northern middle Cenoma-
nian assemblages and other Turonian assemblages from
more southerly WIS environments. This study also gives
insights into euselachian palaeoecology and palaeobiogeog-
raphy within this region of the WIS.
Geological context
Localities
Upper Cretaceous rocks near Watino, Alberta, located at
55◦4250.3 N, 117◦3616.4 W (Fig. 1), provided fossils
that were collected from a sandstone lens along the bank
of the Smoky River that is designated as University of
Alberta Laboratory for Vertebrate Paleontology (UALVP)
locality 76. This locality is at the margin of a large slump
on the east bank of the Smoky River, on a sharp bend
approximately 1 km below the highway bridge. We lack
Figure 1. Map of northwestern Alberta showing the Watino fossil
locality.
detailed information on a second locality, UALVP 848,
from which an earlier collection of euselachian teeth
came. Although collection details are limited, information
associated with the specimens stated, ‘Teeth-Lower Smoky
River-on Smoky River above Watino, T77 R24 W5, at river
level, I.D. Crawford’. All euselachian taxa associated with
this locality are also present at locality 76.
Regional stratigraphical setting
Because of extensive slumping, Cretaceous strata tend to
be poorly exposed in the Smoky River valley. At Watino,
Cretaceous sandstone is exposed at river level and the bone
bed material is known only from loose blocks in heavily
slumped debris. The nearest well-exposed section is at
Hunting Creek, located 8 km NNE of Watino, where the
upper part of the Dunvegan Formation and lower part of the
overlying Kaskapau Formation are exposed (Fig. 2). Using
sea-level as a datum, the top of the Dunvegan Formation
can be traced southward from Erin Lodge on the Peace
River, to Hunting Creek near the Smoky River, and then
projected south to Watino where the Dunvegan-Kaskapau
contact is predicted to lie about 70 m above the level of the
Smoky River. Therefore, rocks exposed near water level in
the vicinity of Watino are probably part of the Dunvegan
Formation, while those above, including the sediments with
the fossil material, are part of the Kaskapau Formation.
An allostratigraphical approach was used to analyse
stratal architecture and facies distribution in the Kaskapau
Formation (Plint 2000; Varban & Plint 2005, 2008a;
Kreitner & Plint 2006). This formation represents a broad,
shallow marine shelf that gradually became less restricted
as a result of progressive marine transgression in the latest
Cenomanian and early Turonian. It contains a number of
units that represent transgressive-regressive events.
Eastward, the Kaskapau strata thin dramatically, from
about 900 m in the Chetwynd area of British Columbia to
<100 m in the Hunting Creek-Watino area of Alberta
(Varban & Plint 2005). In the west, between Chetwynd
and Tumbler Ridge, Kaskapau strata are dominated by
nearshore sandstones, whereas ∼200 km to the east at Erin
Lodge and Hunting Creek, silty claystones and organic
rich calcareous claystones dominate (Varban & Plint
2008a). The eastward thinning of the Kaskapau strata is a
consequence of deposition in an actively subsiding foreland
basin in which stratal packages thin and even lap out
eastwards against the flank of the forebulge (Varban & Plint
2008b). Storm driven geostrophic flows were responsible
for sediment transport across the Kaskapau shelf, which
is interpreted to have been of extremely low relief, always
above storm wave base and probably no more than ∼40 m
deep at a distance of ∼200 km offshore (Varban & Plint
2008a, b). Sand and then silt were progressively extracted
along the transport pathway until only very fine-grained silt
and clay were available at distances of >200 km from shore.
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Turonian Euselachians from Canada 557
Figure 2. Overview photograph of the Hunting Creek section (NTS Grid Reference 83N/13 672852) showing the position of the
Dunvegan-Kaskapau boundary, the regional flooding surfaces X and K1 and the phosphatic sandstone.
Hunting Creek section
Note that the initial interpretation of the Hunting Creek
section (fieldwork in 1993, reported in Plint 2000) has been
revised in the light of subsequent work by Varban (2004),
resulting in several changes including the position of the
top Dunvegan surface being raised by 6 m; the revised
stratigraphical interpretation is shown in Figs 2 and 3.
The Kaskapau Formation exposed at Hunting Creek is
interpreted with reference to well logs to the south and west,
to sections through equivalent strata at Erin Lodge and
Howard Creek (on the Peace River), and to core at 15-34-
77-1W6 (Plint 2000; Varban & Plint 2005). It is important
to recognize that the Kaskapau strata at Hunting Creek are
highly attenuated due to deposition on the flank of the fore-
bulge where subsidence rate was low, and in consequence,
rock units tend to be thin and bounded by unconformities.
Because of limited subsidence, the area was strongly
affected by relative sea-level oscillations that generated a
succession of erosion surfaces and lag deposits.
The lower part of the Kaskapau consists of shallow
marine sandstones and mudstones of the A-X and Doe
Creek units (e.g. Fig. 3; Plint 2000, fig. 13). Above the
K1 surface (a basin-wide disconformity) is a 60 cm bed
of silty clay and 30 cm of fine sandstone; this latter may be
a remnant of the Erin Lodge sandstone that is interpreted
to be of eastern (forebulge) provenance (Varban & Plint
2008a). The Erin Lodge sandstone is sharply overlain (at
52.5 m, Fig. 3) by laminated, slightly silty organic rich
claystone that contains numerous bands and nodules of
fibrous (cone-in-cone) gypsum. This facies is typical of
the ‘Hard Platy Shale’ that crops out extensively along the
Peace River valley (Stelck & Wall 1954; Wallace-Dudley
& Leckie 1995) and which is equivalent to the lower
part of the ‘Second White Speckled Shale’ of subsurface
terminology. On the basis of comparison with the Howard
Creek and Erin Lodge sections, for which biostrati-
graphical control is available (Varban & Plint 2005), the
Cenomanian–Turonian boundary is placed at the base of
the organic rich claystone at 52.2 m (Fig. 3). The claystone
is punctuated at 54.3 m by a 5 cm-thick, quartz-rich,
phosphatic sandstone with sharp lower and upper contacts
(Fig. 3). The detrital grains comprising this bed consist
of about 78% well sorted, angular to subangular, upper
very fine-grained monocrystalline quartz sand, about 2%
glaucony grains and a trace amount of chert and feldspar.
Phosphatic debris forms about 20% of grains and includes
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558 T. D. Cook et al.
Figure 3. Stratigraphical log for the upper part of the section
exposed in Hunting Creek (lower 48 m of Dunvegan Formation
is not shown). Both lithostratigraphical and allostratigraphical
divisions are shown; allostratigraphical interpretations are based
on regional subsurface analyses in Plint (2000) and Varban & Plint
(2005).
bones, scales and teeth. The framework grains are separated
by highly displacive cement that forms about 60% of the
rock volume (Fig. 4A). Cements include calcite, gypsum
and phosphate (Fig. 5A, B); detailed discussion of the
cement paragenesis is beyond the scope of this paper.
A compositionally and texturally similar quartz-rich
phosphatic sandstone, 0–30 cm thick, erosively overlies
pyritic, organic-rich claystone of the Hard Platy Shale at
Howard Creek, 85 km to the west. At this locality, the phos-
phatic sandstone, which is highly lenticular, forms wave
Figure 4. A, micrograph of phosphatic sandstone from Hunting
Creek, showing angular quartz grains and fragmented phosphatic
material in a displacive cement of calcite and minor gypsum;
plane polarized light; B, micrograph of phosphatic sandstone from
Watino, showing angular quartz grains and fragmented phosphatic
material in a displacive cement of calcite and gypsum; plane polar-
ized light.
ripples, hummocky cross-stratification and gutter casts,
suggestive of at least intermittent storm wave action. This
bed can also be traced to the 15-34-77-1W6 core where
it is only 3 cm thick. Although phosphatic lags occur at
other horizons in the claystone facies of the Kaskapau, none
contain more than a trace of quartz sand and therefore the
quartzose bone bed noted at Hunting Creek, Howard Creek
and in the 15-34 core are interpreted to be the same bed.
A micrograph of the phosphatic sandstone from Watino
is shown in Fig. 4B. The detrital grains consist of about
82% well sorted, angular to subangular, very fine-grained
monocrystalline quartz sand, about 3% chert; glaucony
and feldspar are present in trace amounts. Phosphatic
material forms about 15% of the framework grains. The
cement, which forms about 60% of the rock volume, is
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Turonian Euselachians from Canada 559
Figure 5. A, micrograph of phosphatic sandstone from Hunting Creek showing detrital grains and displacive cement; plane polarized
light; B, as A but with crossed nichols showing detrital quartz, chert and phosphatic grains, and calcite and gypsum cement; C, micrograph
of phosphatic sandstone from Watino showing detrital grains and displacive cement; plane polarized light; D, as C with crossed nichols,
showing detrital quartz, chert and phosphate grains, and calcite and gypsum cement.
highly displacive and consists mainly of calcite with minor
gypsum (Fig. 5C, D).
The detrital grains forming the phosphatic sandstones
(Figs 4, 5) bear a very close similarity in terms of grain
size, sorting, angularity and composition, and in the
phosphatic debris. On this basis, it is concluded that the
Hunting Creek phosphatic sandstone and the bone-bearing
sandstone recovered from slumped debris at Watino are
the same bed. Correlation of the phosphatic sandstone bed
from Howard Creek to wells to the south and west suggest
that the bed marks the top of Kaskapau unit II (Varban &
Plint 2005). This bone bed is interpreted as a winnowed
lag deposit and suggests a period of increased wave energy
at the sea floor, an interpretation supported by the presence
of forced-regressive shoreface sandstones at the top of
unit II in the British Columbia Foothills (Varban & Plint
2005). Thus, Kaskapau unit II can be interpreted to have
terminated with a period of relative sea-level fall that led
to forced regression on the western margin of the basin,
and also resulted in shallowing and erosion of offshore
claystones in the distal, forebulge region, accompanied by
concentration of phosphatic debris.
Materials and methods
Most specimens were surface collected, although others
were obtained from acid dissolved matrix. Larger teeth
embedded in the sandstone matrix were mechanically
removed using a pneumatic hand drill and a dental pick.
Teeth deeply embedded within matrix were placed in a
buffered 10% acetic acid solution, which was effective
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560 T. D. Cook et al.
in dissolving the calcite cement binding the sandstone.
Following the acid bath, the specimens were washed
with water to remove any acid residue. Specimens were
separated from the dissolved matrix using a 0.5 mm sieve,
followed by manual picking of teeth from the residues. A
binocular microscope was used to examine and identify
the isolated euselachian teeth. Any fractured teeth were
hand picked and repaired using an adhesive. For detailed
imaging of small teeth, a Jeol Field Emission Scanning
Electron Microscope (JSM-6301 FXV) was used. Larger
teeth were coated with ammonium chloride and imaged
using a Nikon DXM 1200c digital camera mounted on a
Zeiss Discovery V8 stereo microscope.
The Watino teeth are catalogued in the collections
of the University of Alberta Laboratory of Vertebrate
Paleontology (UALVP).
Tooth morphological terminology largely follows
Cappetta (1987), whereas tooth type terminology includes
information presented by Siverson (1999) and Shimada
(2002).
Systematic palaeontology
Class Chondrichthyes Huxley, 1880
Subclass Elasmobranchii Bonaparte, 1838
Cohort Euselachii Hay, 1902
Order Hybodontiformes Maisey, 1989
Fam ily Hybodontidae Owen, 1845
Genus Meristodonoides Underwood & Cumbaa, 2010
Meristodonoides cf. rajkovichi (Case, 2001)
(Fig. 6A, B)
Material. UALVP 53121, fractured tooth, locality 76.
UALVP 53124, fractured tooth, locality 76. UALVP 53123,
fractured tooth, locality 848.
Description. These specimens are median cusps of highly
fractured teeth. The apex of each cusp is missing. The
strongly convex lingual and labial faces contain numerous
large, widely spaced, longitudinal ridges. The labial ridges
are restricted to the lower third of the cusp, whereas the
lingual ridges extend two thirds up the cusp. A distinct
cutting edge extends the full length of the medial cusp.
The cusp has a lingual curvature in profile. The teeth are
missing the root.
Remarks. Underwood & Cumbaa (2010) erected the
genus Meristodonoides to accommodate the teeth of certain
species previously ascribed to Hybodus Agassiz, 1837. The
included taxa are Meristodonoides rajkovichi,M. butleri
(Thurmond, 1971), M. montanensis (Case, 1978) and M.
novojerseyensis (Case & Cappetta, 2004). According to
Underwood & Cumbaa (2010, p. 906), the teeth of these
species can be differentiated from the teeth of species of
Hybodus by “the presence of a single well-developed cusp,
very low root and, for some species, lack of a labial boss”.
Species of Meristodonoides recovered from northern
localities in Canada include M. rajkovichi,M. butleri
and M. montanensis. Numerous M. rajkovichi teeth were
reported by Cumbaa et al. (2006; as Hybodus sp.) and
Underwood & Cumbaa (2010) from the Bainbridge bone
bed in east central Saskatchewan. A small number of teeth
were assigned to M. butleri by Cumbaa et al. (2006; as
H. butleri). Underwood & Cumbaa (2010) suggested that
the tooth figured by Cook et al. (2008, fig. 4a) as Hybodus
sp. from the middle Cenomanian Dunvegan Formation
of Alberta should be assigned to M. rajkovichi.Givenits
fragmentary condition, Cook et al. (2008) conservatively
left the reported specimen in open nomenclature; however,
the crown contour and ornamentation of the specimen does
resemble teeth figured as M. rajkovichi (see Case 2001,
pl. 1, fig. 5, pl. 2, figs 1, 2; Underwood & Cumbaa 2010,
pl. 1, figs 3–17).
Meristodonoides butleri, originally reported from the
Aptian–Albian of Texas (Thurmond 1971; as Hybodus
butleri), differs from M. rajkovichi by possessing a shorter
and broader medial cusp with stronger ornamentation
(Underwood & Cumbaa 2010). Meristodonoides monta-
nensis has been reported (as H. montanensis)fromthe
late Campanian of Montana (Case 1978), Wyoming (Case
1987) and Alberta (Beavan & Russell 1999). The teeth of
this species are characterised by their medium to large size
and vertical enameloid folds that extend only one-quarter
the height of the medial cusp (Case 1978). Due to the incom-
plete nature of specimens UALVP 53121 and 53122, species
determination is problematic. However, based strictly on
cusp morphology, the specimens most closely resemble M.
rajkovichi. Despite this conservative identification, it should
be noted that Meristodonoides material at the Watino local-
ities is rare, with only three specimens collected. This is in
stark contrast to the abundance of Meristodonoides teeth
reported in the middle Cenomanian Saskatchewan assem-
blage (Cumbaa et al. 2006; Underwood & Cumbaa 2010).
Fam ily Polyacrodontidae Glickman, 1964
Genus Polyacrodus Jaekel, 1889
Polyacrodus sp.
(Fig. 6C)
Material. UALVP 53124, fractured tooth, locality 76.
Description. This specimen consists of a highly fractured
crown containing a blunt median cusp that is very slightly
distally inclined. Distal to the cusp are two lateral cusplets;
the more distal cusplet is significantly reduced. The portion
of the tooth mesial to the median cusp is fractured. The
lingual and labial crown faces are strongly convex and
contain multiple well developed enameloid folds that
extend from the crown foot to the apex of the medial cusp
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Turonian Euselachians from Canada 561
Figure 6. Meristodonoides,Polyacrodus and Ptychodus teeth from the Watino localities. A,B, Meristodonoides cf. rajkovichi (Case,
2001); A, fractured tooth, UALVP 53121; B, fractured tooth, UALVP 53122; C, Polyacrodus sp., fractured tooth, UALVP 53124; D–H,
Ptychodus anonymus Williston, 1900; D, medial tooth, UALVP 53125; E, medial tooth, UALVP 53126; F, lateral tooth, UALVP 53127;
G, lateral tooth, UALVP 53128; H, lateral tooth, UALVP 53129. Views: labial (left), profile (centre), and lingual (right) for A; labial (left)
and lingual (right) for B; labial (top) and occlusal (bottom) for C; occlusal (left) and profile (right) for D and E; occlusal for F–H. A, C,
D, F, and G are SEM images. Scale bars =1 mm.
and lateral cusplets. A weak labial protuberance is present.
The root is missing.
Remarks. Isolated teeth belonging to this genus have been
reported from the Lower Triassic to the Upper Cretaceous
of Europe and Greenland (Cappetta 1987). Given the poor
preservation of UALVP 53124, we cautiously assign this
tooth to Polyacrodus sp. based on the low median cusp
and lateral cusplets, weak labial protuberance, and strong
enameloid folding that extends to the crown apex.
Teeth of Polyacrodus have been recovered from other
Turonian localities within the WIS, including the middle
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562 T. D. Cook et al.
Turonian of Texas (Welton & Farish 1993; as Polyacrodus
illingsworthi (Dixon, 1850)) and late Turonian of South
Dakota and Wyoming (Cappetta, 1973; as Synechodus
sp.; Evetts 1979, as S.illingsworthi; Cicimurri 2004;
as Polyacrodus illingsworthi). UALVP 53124 is the first
documented occurrence of Polyacrodus in Canada and
expands its known northern geographical range within
the WIS. Rees & Underwood (2002, 2008) questioned
the validity of this genus as the tooth morphology closely
resembles that of certain Hybodus species. Unfortunately,
the incomplete nature of UALVP 53124 does not allow us
to contribute any additional insights into this matter.
Order Incertae ordinis
Fam ily Ptychodontidae Jaekel, 1898
Genus Ptychodus Agassiz, 1835
Ptychodus anonymus Williston 1900
(Fig. 6D–H)
Material. UALVP 53125, medial tooth, locality 76.
UALVP 53126, medial tooth, locality 848. UALVP 53127,
lateral tooth, locality 76. UALVP 53128, lateral tooth,
locality 848. UALVP 53129, lateral tooth, locality 848.
UALVP 53130, nine teeth, locality 848.
Description. Teeth positioned medially in the jaw possess
a crown with a high and rounded cusp. Restricted to
this cusp are multiple fine transverse ridges that do not
extend into the surrounding granular marginal region.
The crown overhangs the root in all directions and has a
distinct concave notch along its lingual margin. The root
is positioned somewhat lingually under the crown and
contains multiple foramina of various sizes with larger
foramina situated under the crown-root border. The labial
root face is flat and is at an oblique angle to the base of
the crown. The lingual root face may be slightly convex or
concave. The basal root face is more or less flat.
Laterally positioned teeth contain a low asymmetrical
crown that overhangs the root in all directions. There is a
small rounded cusp placed lingually on the crown. Distinct
transverse ridges extend across the cusp, with many of
them bifurcating upon reaching the reduced granular
margin. The root is low and bears multiple foramina.
Larger foramina are located near the crown-root boundary.
Remarks. TheteethofPtychodus are arranged in a
row-locking configuration that forms a crushing pavement
dentition in both jaws (see Shimada et al. 2009). The
symmetrical nature of UALVP 53125 and 53126 suggests
that these teeth are from a medial position, whereas the
more asymmetrical UALVP 53127, 53128 and 53129
are from a more lateral position. The latter teeth bear
some resemblance to teeth of Ptychodus rhombodus, first
described from the Bainbridge bone bed of Saskatchewan
by Underwood & Cumbaa (2010). According to these
authors, the small teeth of P. rhombodus are unique among
other described species of Ptychodus in possessing a low
and flat crown that bears a small but distinct cusp and lacks
a defined marginal region. The teeth of P. rhombodus can
be distinguished from P. anonymus by the presence of a
more developed cusp and distinct granular margin in the
latter (Underwood & Cumbaa 2010); these features are
observed in the specimens described herein.
Many Ptychodus anonymus teeth, recovered from
various assemblages throughout the WIS, have been re-
diagnosed recently as P. rugosus Dixon, 1850. According to
Hamm (2010a), distinct crown morphologies separate these
two species. In addition, “Ptychodus anonymus is found
in Middle Cenomanian through Middle Turonian deposits,
whereas P. rugosus is known only from Late Coniacian
through Late Santonian deposits” (Hamm 2010a, p. 45).
In accordance with this re-assessment, P. anonymus has
been recovered from the late Cenomanian and Turonian
of Texas (Welton & Farish 1993), the middle Turonian of
New Mexico (Wolberg 1985), the middle Cenomanian
of Colorado (Shimada et al. 2006), the middle Turonian of
Kansas (Fielitz & Shimada 1999), the late Cenomanian and
middle Turonian of South Dakota and Wyoming (Cicimurri
2001a, 2004) and the early Turonian of Saskatchewan
(Case et al. 1990; as Ptychodus cf. P. rugosus). A medial
tooth of P. anonymus was reported from the Saskatchewan
Bainbridge bone bed by Cumbaa et al. (2006); however,
Underwood & Cumbaa (2010) subsequently reassigned this
tooth to their new species P. rhombodus. Also recovered
from the Saskatchewan bone bed were two teeth indentified
as Ptychodus sp. that resembled P. anonymus but lacked the
granular marginal area (Underwood & Cumbaa 2010). The
tooth morphology of P. anonymus also closely resembles
P. mammillaris Agassiz, 1835; however, the latter has fewer
and coarser transverse ridges and a concentric granular
pattern on the marginal region. The specimens herein
represent the first description of P. anonymus from Alberta
and extend the northern geographical range of this species.
Order Lamniformes Berg, 1958
Fam ily Anacoracidae Casier, 1947
Genus Squalicorax Whitley, 1939
Squalicorax sp. A
(Fig. 7A–M)
Material. UALVP 53131, anterior tooth, locality 76.
UALVP 53141, anterior tooth, locality 848. UALVP 53142,
symphyseal tooth, locality 76. UALVP 53143, lateral tooth,
locality 848. UALVP 53144, lateral tooth, locality 848.
UALVP 53145, lateral tooth, locality 76. UALVP 53146,
lateral tooth, locality 76. UALVP 53147, lateral tooth,
locality 76. UALVP 53148, lateral tooth, locality 848.
UALVP 53149, lateral tooth, locality 76. UALVP 53150,
lateral tooth, locality 848. UALVP 53151, lateral tooth,
locality 76. UALVP 53152, fractured lateral tooth, locality
76. UALVP 53153, 19 complete and fractured teeth, locality
76. UALVP 53154, 95 complete and fractured teeth, locality
848.
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Turonian Euselachians from Canada 563
Figure 7. Squalicorax sp. A teeth from the Watino localities; A, anterior tooth, UALVP 53131; B, lower anterior tooth, UALVP 53141;
C, symphyseal tooth, UALVP 53142; D, lateral tooth, UALVP 53143; E, lateral tooth, UALVP 53144; F, lateral tooth, UALVP 53145; G,
lateral tooth, UALVP 53146; H, lateral tooth, UALVP 53147; I, lateral tooth, UALVP 53148; J, lateral tooth, UALVP 53149; K, lateral
tooth, UALVP 53150; L, lateral tooth, UALVP 53151; M, fractured lateral tooth, UALVP 53152. Note that the root of B and G is slightly
eroded. Views: labial (left), profile (centre), and lingual (right) for A; labial (top) and lingual (bottom) for C; labial (left) and lingual (right)
for B and D–M. D is an SEM image. Scale bars =1 mm.
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564 T. D. Cook et al.
Description. The upper anterior teeth are taller than wide
and have a cusp that is slightly distally inclined. The labial
crown face is slightly convex, whereas the lingual crown
face is strongly convex. The mesial cutting edge is slightly
sigmoidal and contains coarse serrations that become
reduced in size towards the apex. This cutting edge forms a
sharp apex with the distal cutting edge, which is shorter and
also contains serrations that become more robust towards
the base. The distal heel is short, low and finely serrated.
There is a distinct lingual neck that is slightly wider in
the medial region of the crown. The sigmoidal labial basal
ledge of the crown overhangs the labial root face, which
is particularly prominent in the medial region. The distal
root lobe is slightly more robust than the mesial root lobe.
These rounded root lobes are separated by a distinct basal
concavity. Multiple small foramina are present throughout
the labial, lingual and basal root faces with larger foramina
located at the apex of the lingual protuberance and below
the labial crown-root border. Lower anterior teeth have a
more erect and narrow cusp.
Symphyseal teeth are considerably smaller and mesodis-
tally compressed. The labial crown face is flat, whereas
the lingual crown face is convex. The mesial cutting edge
is irregular and sigmoidal, whereas the distal cutting edge
is slightly convex and smooth. A distinct distal heel also
lacks serrations. The distal lobe of the root is more robust
than the mesial lobe and the basal concavity is shallow.
Teeth from a more lateral position are wider than tall and
possess a cusp that is strongly distally inclined. Typically,
the basal one-third of the mesial cutting edge is straight,
whereas the remainder of the cutting edge is slightly
convex. The distal cutting edge is shorter and is either
straight or slightly convex. Both cutting edges have well
developed serrations that become smaller towards the apex.
The heel is straight to convex and bears fine serrations.
The height of the lingual neck is relatively constant across
the tooth. The sigmoidal labial basal ledge of the crown
is well developed and overhangs the labial root face. The
distal root lobe is slightly more robust than the mesial root
lobe and the basal concavity may be quite shallow in the
most distally positioned teeth.
Remarks. Teeth from many North American assemblages
that were initially reported as Squalicorax falcatus (Agas-
siz, 1843) (e.g. Case et al. 1990; Williamson et al. 1993;
Cumbaa et al. 2006; Shimada et al. 2006) have been
reassigned to S. curvatus (Williston, 1900) by Underwood
& Cumbaa (2010). This corroborates the notion presented
by Siverson et al. (2007) that teeth identified as S. falcatus
by some North American authors do not agree with the
lectotype from the Turonian chalk of England and that the
presence of S. falcatus within North American deposits is
suspect.
The teeth described herein conform to the general
morphology of Squalicorax curvatus, originally described
and illustrated by Williston (1900, pl. 30, fig. 8), as Corax
curvatus, from the Benton Formation of Kansas. Siverson
et al. (2007) questioned the validity of this species, as
the description was based on only two poorly preserved
syntypes. We agree with the concerns of Siverson et al.
(2007) about the validity of S. curvatus and also regard it as a
nomen dubium pending re-examination of the type material
and recovery of additional topotypes. Despite this issue, the
teeth listed here have a similar overall morphology to teeth
described as Squalicorax sp. from the Dunvegan Formation
of Alberta (Cook et al. 2008) and S. curvatus from the
Bainbridge bone bed and Favel Formation of Saskatchewan
(Case et al. 1990; Underwood & Cumbaa 2010).
Squalicorax sp. B
(Fig. 8A–M)
Material. UALVP 53155, symphyseal tooth, locality
76. UALVP 53156, anterior tooth, locality 76. UALVP
53157, anterior tooth, locality 76. UALVP 53158, fractured
anterior tooth, locality 848. UALVP 53159, lateral tooth,
locality 848. UALVP 53160, lateral tooth, locality 848.
UALVP 53161, lateral tooth, locality 76. UALVP 53162,
lateral tooth, locality 848. UALVP 53163, lateral tooth,
locality 76. UALVP 53164, lateral tooth, locality 848.
UALVP 53165, lateral tooth, locality 848. UALVP 53166,
lateral tooth, locality 76. UALVP 53167, lateral tooth,
locality 76. UALVP 53168, 17 complete and fractured
teeth, locality 76. UALVP 53169, 158 complete and
fractured teeth, locality 848. UALVP 53170, 206 complete
and fractured teeth, locality 848.
Description. Anterior teeth are taller than wide. The crown
contains a median cusp that is slightly distally inclined. The
upper two-thirds of the labial crown face are slightly convex,
whereas the basal one-third is flat. The lingual crown face is
strongly convex. Both faces are smooth. The mesial cutting
edge is gently sigmoidal and is devoid ofdistinct serrations.
The apex forms an acute angle. The distal cutting edge is
straight and shorter than the mesial cutting edge and also
lacks serrations. It intersects the smooth distal heel at an
obtuse angle. The lingual neck is broad, particularly in
the medial region. The labial basal ledge is sigmoidal and
overhangs the labial root face to a greater extent medially.
The bilobate root contains a weak lingual root protuberance
and lacks a nutrient groove. The distal root lobe is typically
more robust than the mesial root lobe. These rounded root
lobes are separated by a deep basal concavity. The root is
porous with small foramina present throughout the labial,
lingual and basal root faces. Larger foramina are typically
located under the labial crown-root boundary.
Symphyseal teeth are significantly smaller and mesodis-
tally compressed. The labial crown face is flat, whereas the
lingual crown face is convex. The cutting edges are smooth
and there is a distinct distal heel. The root has a lingual
protuberance and is porous, containing multiple foramina.
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Turonian Euselachians from Canada 565
Figure 8. Squalicorax sp. B teeth from the Watino localities. A, anterior tooth, UALVP 53156; B, lateral tooth, UALVP 53157; C,
symphyseal tooth, UALVP 53155; D, fractured lateral tooth, UALVP 53158; E, lateral tooth, UALVP 53159; F, lateral tooth, UALVP
53160; G, lateral tooth, UALVP 53161; H, lateral tooth, UALVP 53162; I, lateral tooth, UALVP 53163; J, lateral tooth, UALVP 53164;
K, lateral tooth, UALVP 53165; L, lateral tooth, UALVP 53166; M, lateral tooth, UALVP 53167. Note that the root of A, B, F, and M
is slightly eroded. Views: labial (left), profile (centre), and lingual (right) for A; labial (top) and lingual (bottom) for C; labial (left) and
lingual (right) for B and D–M. G is an SEM image. Scale bars =1 mm.
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566 T. D. Cook et al.
A very shallow basal concavity separates the mesial root
lobe from the smaller distal root lobe.
Lateral teeth are wider than tall and possess a narrow
median cusp that is strongly distally inclined. Both the
mesial and distal cutting edges are straight to slightly
convex. Both edges lack distinct serrations but weak
crenulations may be present. A distinct notch may be
present where the distal cutting edge and the distal heel
meet. The distal heel lacks serrations. A lingual neck is
broad and may bear short ridges. The labial basal crown
edge is sigmoidal and overhangs the labial root face. In
the distal-most files, this ledge is more or less straight. The
root is porous. The distal root lobe is more robust than
the mesial root lobe. The deep basal concavity becomes
shallow in only the most distally positioned teeth.
Remarks. The teeth of Squalicorax sp. B can be distin-
guished from Squalicorax sp. A by possessing: (1) a
median cusp that is narrower; (2) a straighter mesial cutting
edge; (3) smooth cutting edges; (4) a higher distal heel;
(5) a more developed distal notch; (6) a distal root lobe
more robust than the mesial root lobe; (7) a more porous
root; and (8) a deeper basal concavity on all but the most
distally positioned teeth. Although Squalicorax sp. A teeth
are abundant at Watino, the teeth of Squalicorax sp. B are
three times more common.
The tooth morphology of Squalicorax sp. B is indistin-
guishable from that of a species recovered from the Haycock
Marl of Western Australia (Siverson 1996; Siverson et al.
2007) and we consider them conspecific. A re-description
of the Haycock Marl specimens is currently being under-
taken (M. Siverson pers. comm. 2010). The Australian teeth
(originally described as S. volgensis (Glickman in Glick-
man & Shvazhaite, 1971) from the Beedagong Claystone in
Siverson 1996) and the Watino teeth are similar in overall
size and have narrow cusps that lack distinct serrations.
Other comparable features include similar lingual neck and
labial basal crown ledge morphology, a high distal heel, and
a distal root lobe that is considerably more robust than the
mesial lobe and is separated by a very deep basal concavity.
Squalicorax sp. B can be distinguished from teeth
identified as S. pawpawensis Siverson et al., 2007 from
the late Albian of Texas, which possess more developed
serrations along the cutting edges, an almost straight labial
basal crown ledge and a shallower basal concavity sepa-
rating the root lobes. Teeth recovered from the Bainbridge
of Saskatchewan and identified as Paleoanacorax aff.
pawpawensis by Underwood & Cumbaa (2010, p. 935), are
smaller and contain “faint crenulations or incipient serra-
tions” on the mesial cutting edge of most teeth. In addition,
the root is more robust and the basal concavity is shallower.
The overall morphology of the Saskatchewan teeth is differ-
ent from the Texan Squalicorax pawpawensis and likely
represents a new species. Squalicorax sp. B is also distinct
from the teeth identified as S. volgensis from the early
Cenomanian of Russia (Glickman & Shvazhaite 1971).
The latter has a more prominent overhang of the labial
basal crown ledge and cutting edges that are very weakly
serrated (see Siverson et al. 2007, text-fig. 4c–f). Teeth
identified as S. volgensis from the middle Cenomanian of
Saskatchewan (Cumbaa et al. 2006, fig. 4.7) and South
Dakota (Cicimurri 2001b, fig. 5h), the late Cenomanian of
Kansas (Shimada & Martin 2008, fig. 5c), and the Turonian-
Coniacian boundary of Texas (Cappetta & Case 1999,
pl. 5, fig. 1) conform to the tooth morphology of Squalico-
rax sp. B herein, and to the Haycock Marl specimens.
Fam ily Archaeolamnidae Underwood & Cumbaa, 2010
Genus Archaeolamna Siverson, 1992
Archaeolamna ex. gr. kopingensis (Davis, 1890)
(Fig. 9A–F)
Material. UALVP 53171, lateral tooth, locality 76.
UALVP 53172, anterior tooth, locality 76. UALVP 53173,
fractured anterior tooth, locality 848. UALVP 53174,
fractured anterior tooth, locality 76. UALVP 53176, lateral
tooth, locality 76. UALVP 53177, lateral tooth, locality
848. UALVP 53178, lateral tooth, locality 76. UALVP
53179, 32 fractured teeth, locality 76. UALVP 53180, 45
fractured teeth, locality 848.
Description. Anterior teeth contain a tall, broad-based
triangular median cusp that is erect; however, the third
upper anterior teeth (=intrabullar intermediate teeth of
Shimada 2002) have a median cusp in which the apical
half is strongly distally curved (see Cook et al. 2011).
The labial face of the crown is slightly convex, whereas
the lingual face is strongly convex. Both crown faces
are smooth. A pair of triangular lateral cusplets flanks
the median cusp. A sharp cutting edge runs continuously
across the median cusp and the lateral cusplets. The height
of the narrow lingual neck is relatively constant between
the lateral cusplet and the median cusp regions. The root is
robust and has a well-developed lingual protuberance with
one or more nutrient foramina. A large portion of the root
lobes are missing in the specimens recovered from Watino.
Lateral teeth have a median cusp that is strongly distally
inclined. The smooth labial crown face is slightly convex,
whereas the smooth lingual crown face is strongly convex.
The cutting edge runs continuously between the median
cusp and a pair of triangular lateral cusplets. A narrow
lingual neck is present. The root contains a well-developed
lingual protuberance with a large foramen that may or may
not be situated in a shallow nutrient groove. The root lobes
are more or less symmetrical and separated by a deep basal
concavity that becomes shallow in the distal-most lateral
teeth.
Remarks. The teeth of Archaeolamna have been recov-
ered from numerous Albian to Maastrichtian deposits
throughout the WIS, Europe and Australia, as summarized
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Turonian Euselachians from Canada 567
Figure 9. Archaeolamna and Cardabiodon teeth from the Watino localities. A–F, Archaeolamna ex. gr. kopingensis (Davis, 1890); A,
lateral tooth, UALVP 53171; B, anterior tooth, UALVP 53172; C, fractured anterior tooth, UALVP 53174; D, lateral tooth, UALVP 53176;
E, lateral tooth, UALVP 53177; F, lateral tooth, UALVP 53178; G–I, Cardabiodon aff. C. ricki (Siverson, 1999); G, fractured juvenile
anterior tooth, UALVP 53175; H, lateral tooth, UALVP 49430; I, small juvenile anterior tooth, UALVP 53181. Views: labial (left), profile
(centre), and lingual (right) for A and H; labial (left) and lingual (right) for B–E and G; labial (top) and lingual (bottom) for F; labial (top),
profile (center), and lingual (bottom) for I. B and I are SEM images. Scale bars for A–G and I =1 mm; scale bar for H =1cm.
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568 T. D. Cook et al.
by Cook et al. (2011). Subtle differences in tooth morphol-
ogy from different stratigraphical horizons suggest that
this taxon is likely to represent multiple species (Cumbaa
et al. 2006; Cook et al. 2008; Underwood & Cumbaa
2010), although only two species have been named to
date. The teeth herein share a similar morphology to
middle Cenomanian teeth recovered from the Bainbridge
bone bed assemblage of Saskatchewan and described as
Archaeolamna ex. gr. kopingensis (Underwood & Cumbaa
2010) and we follow this taxonomic assignment. Similar
teeth were also recovered from the middle Cenomanian
Dunvegan assemblage of Alberta (Cook et al. 2008).
However, we note that the teeth of Archaeolamna are also
very similar to juvenile Dwardius Siverson, 1999 (Siverson
1999; M. Siverson pers. comm. 2011) and it is possible that
many of the teeth recovered from Watino may belong to this
latter taxon instead. In addition, specific identification is
particularly problematic due to the eroded nature of many
of the recovered teeth. Therefore, we here retain these teeth
in Archaeolamna on the basis on their similarity to the
material from Saskatchewan but we recognise that future
study may well show them to belong to another taxon.
Fam ily Cardabiodontidae Siverson, 1999
Genus Cardabiodon Siverson, 1999
Cardabiodon aff. C. ricki Siverson, 1999
(Fig. 9G–I)
Material. UALVP 49430, lateral tooth, locality 848.
UALVP 53181, small juvenile anterior tooth, locality 848.
UALVP 53175, juvenile fractured lateral tooth, locality 76.
Description. The lateral tooth contains a large median
cusp that is distally inclined. The labial face is convex.
The labial basal ledge of the crown overhangs the labial
root face. The lingual face is strongly convex and contains
a distinct lingual neck that is thicker medially. A distinct
cutting edge runs continuously between the median cusp
and a pair of blunt lateral cusplets. A pair of weakly
developed heels occur lateral to the cusplets. The large
bilobate root has a large lingual protuberance with no
nutrient groove. The root lobes are asymmetrical, with
the mesial lobe being slightly longer and narrower than
the more robust distal lobe. Numerous foramina are
distributed throughout the labial face of the root, with a
noticeable concentration situated at the crown-root border.
The U-shaped basal concavity is deep.
The small juvenile anterior tooth has a crown with a trian-
gular robust median cusp that is slightly distally inclined
and lingually directed. The slightly convex labial face and
strongly convex lingual face are smooth. The distal cusplet
is missing, whereas the short triangular medial cusplet is
directed distally. The cutting edge is continuous between the
median cusp and the cusplet. A distinct broad lingual neck
is present. The labial basal ledge of the crown overhangs
the labial root face. The bilobate root is missing the medial
root lobe. The lingual protuberance is high, massive, and
contains a large nutrient foramen. Numerous smaller foram-
ina are distributed throughout the root faces but are partic-
ularly concentrated below the labial crown root border.
The juvenile lateral tooth has a median cusp with a
strong distal curvature that is flanked by a pair of well-
developed lateral cusplets. The labial crown face is weakly
convex, whereas the lingual crown face is strongly convex.
The distinct lingual neck is much wider in the medial
region of the crown. The labial basal ledge of the crown
slightly overhangs the labial root face. The root contains
a somewhat robust lingual protuberance. The root lobes
are elongated and the distal portion of the distal root lobe
is missing. Numerous large foramina are predominantly
concentrated below the labial crown root border.
Remarks. Cook et al. (2010) reported the first North
American occurrence of Cardabiodon ricki based on
UALVP 49430 recovered from the UALVP locality 848. In
that document, the Watino locality was described as being
late Cenomanian in age; however, subsequent work (see
above) now indicates an early Turonian age for the Watino
localities.
UALVP 53181 is the first description of a small juvenile
tooth from this species. Juvenile teeth of the congeneric,
younger species Cardabiodon venator Siverson & Lind-
gren, 2005 have a similar overall morphology but a less
developed labial basal ledge overhang than that of C. ricki
(M. Siverson pers. comm. 2010). UALVP 53175 represents
a lateral tooth from an older juvenile. The tooth has a
similar morphology to a lateral tooth of C. venator figured
by Siverson & Lindgren (2005, fig. 3g) but contains
well-developed lateral cusplets.
The various deposits from which the remains of
Cardabiodon ricki have been recovered are summarized
in Cook et al. (2010). As discussed in that document,
UAVLP 49430 has the tall cusp and enlarged lateral
cusplets of C. ricki from the middle Cenomanian but has
a foraminal pattern resembling C. venator from the middle
Turonian. Consequently, the early Turonian teeth described
herein may represent a transitional form. As such, we
now conservatively identify this material as Cardabiodon
aff. C. ricki. A tooth described as Cretolamna woodwardi
Herman, 1977 by Williamson et al. (1993) from the basal
Turonian of Arizona (J. I. Kirkland pers. comm. 2011) was
reassigned to Cardabiodon by Siverson & Lindgren (2005).
Fam ily Cretoxyrhinidae Glickman, 1958
Genus Cretoxyrhina Glickman, 1958
Cretoxyrhina mantelli (Agassiz, 1843)
(Fig. 10A–H)
Material. UALVP 53183, fractured anterior tooth, locality
76. UALVP 53184, fractured lateral tooth, locality 76.
UALVP 53185, fractured lateral tooth, locality 76. UALVP
53186, lateral tooth, locality 76. UALVP 53187, lateral
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Turonian Euselachians from Canada 569
Figure 10. Cretoxyrhina mantelli (Agassiz 1843) teeth from the Watino localities. A, fractured anterior tooth, UALVP 53183; B, lateral
tooth, UALVP 53186; C, fractured lateral tooth, UALVP 53185; D, fractured lateral tooth, UALVP 53184; E, lateral tooth, UALVP 53187;
F, lateral tooth, UALVP 53188; G, lateral tooth, UALVP 53189; H, lateral tooth, UALVP 53190. Views: labial (left), profile (centre), and
lingual (right) for A; labial (left) and lingual (right) for B–H. Scale bars =5 mm.
tooth, locality 76. UALVP 53188, lateral tooth, locality
848. UALVP 53189, lateral tooth, locality 76. UALVP
53190, lateral tooth, locality 848. UALVP 53191, 11
fractured teeth, locality 76. UALVP 53192, 14 fractured
teeth, locality 848.
Description. Anterior teeth are relatively large and contain
a narrow median cusp that is erect or slightly distally
inclined depending upon position in the jaw. The flat labial
and strongly convex lingual faces are both smooth and
contain a cutting edge that is continuous to the crown foot
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570 T. D. Cook et al.
and sigmoidal in profile view. Lateral cusplets are absent.
The lingual neck is broad medially. The mesial root lobe
is slightly more elongated and is separated from the distal
root lobe by a deep basal concavity. A well-developed
lingual protuberance is present and contains a large nutrient
foramen but lacks a nutrient groove. Additional foramina
are present throughout the labial root face.
Lateral teeth possess a broad and distally inclined
median cusp that is flanked by a pair of short and broad
lateral cusplets on most teeth. A cutting edge runs across
these cusps. The lingual neck is narrow. The root lobes
are more or less symmetrical and become more divergent,
thus forming a shallow basal concavity in the most distal
lateral teeth. The lingual protuberance is well developed
and contains a large nutrient foramen. Additional foramina
are present throughout the labial root face.
Remarks. By comparing the maximum tooth size of
the Watino specimens with Cretoxyrhina teeth recovered
from other Cenomanian and Turonian WIS assemblages
(Welton & Farish 1993; Williamson et al. 1993; Siverson
& Lindgren 2005; Shimada et al. 2006; Shimada & Martin
2008; Underwood & Cumbaa 2010), it is evident that the
recovered teeth are from juvenile and sub-adult forms.
Siverson & Lindgren (2005) noted that Cretoxyrhina was
in the process of cusplet reduction during the late Albian
to middle Turonian. Lateral cusplets in teeth identified
as C. mantelli by Siverson & Lindgren (2005), from the
lower middle Turonian of Montana, are restricted to only
the distally situated lateral teeth in both juvenile and sub-
adult/adult forms. The Watino specimens likely represent
a transitional form of Cretoxyrhina, in which the lateral
cusplets of anterior teeth are lost in both juvenile and sub-
adult stages (and presumably adult forms) but are retained
in most juvenile and sub-adult lateral teeth although
reduced. Juvenile anterior teeth from the late Cenomanian
of Kansas (Shimada & Martin 2008, fig. 6e) also lack
lateral cusplets and share similar general tooth morphology.
The overall morphology of the teeth described herein also
closely resembles Cretoxyrhina material, identified as C.
denticulata (Glickman, 1957), from the middle Cenoma-
nian Bainbridge bone bed of Saskatchewan (Underwood
& Cumbaa 2010). Unlike the Watino teeth, most anterior
teeth from Saskatchewan “have a pair of incipient to very
low lateral cusplets” (Underwood & Cumbaa 2010, p. 912).
Lateral teeth assigned to Cretodus sp. by Case et al. (1990,
fig. 7a–d) from the Favel Formation of Saskatchewan likely
belong to small juvenile Cretoxyrhina mantelli.
Fam ily Mitsukurinidae Jordan, 1898
Genus Scapanorhynchus Woodward, 1889
Scapanorhynchus sp.
(Fig. 11A)
Material. UALVP 53193, fractured lateral tooth, locality
76.
Description. The large incomplete lateral tooth contains
a tall, broad-based median cusp that is labiolingually
compressed and distally inclined. The smooth labial crown
face is flat, whereas the basal one-third of the weakly
convex lingual crown face bears strong parallel striations.
A distinct cutting edge runs the entire length of the cusp
and is sigmoidal in profile view. A narrow lingual neck is
present. The preserved portion of the root is very low and
contains a strong lingual protuberance that bears a deep
nutrient groove. The root lobes are missing but are typically
sub-rectangular in specimens recovered from other deposits
(see Siverson 1992, pl. 4, figs 8–11; Welton & Farish 1993,
p. 94, figs 1, 4; Cappetta & Case 1999, pl. 11, figs 8–11).
Remarks. The labiolingually compressed and sigmoidal
median cusp, the low medial region of the root and the paral-
lel basal striations of UALVP 53193 are characters observed
in the lateral teeth of Scapanorhynchus. The latter feature
distinguishes the teeth of this taxon from the irregular and
flexuous striations of odontaspidid teeth (Siverson 1992).
Scapanorhynchus raphiodon (Agassiz, 1843) has been
reported from Cenomanian–Coniacian deposits throughout
the WIS (e.g. Wolberg 1985; Welton & Farish 1993;
Williamson et al. 1993; Cicimurri 2001a, 2004; Hamm
& Shimada 2002; Becker et al. 2010). Siverson (1992)
questioned the validity of this species. Unfortunately, the
incomplete nature of this specimen precludes determination
to species level and further taxonomic discussion.
Fam ily Odontaspididae M¨
uller & Henle, 1839
Genus Carcharias Rafinesque, 1810
Carcharias aff. C. striatula (Dalinkevicius, 1935)
(Fig. 11B–E)
Material. UALVP 53194, fractured anterior tooth, locality
76. UALVP 53195, lateral tooth, locality 848. UALVP
53196, lateral tooth, locality 76. UALVP 53197, lateral
tooth, locality 848.
Description. The anterior tooth possesses an erect broad-
based median cusp that tapers to a sharp apex and is slightly
sigmoidal in profile view. The labial crown face is weakly
convex and strongly overhangs the labial root face. This
basal ledge may be smooth or bear weak striations that are
restricted to the ledge. The lingual crown face is strongly
convex and bears weak flexuous striations that are restricted
to the basal half of the median cusp. The cutting edges of the
median cusp do not extend to the pair of short needle-like
lateral cusplets. A narrow lingual neck separates the median
cusp and lateral cusplets from the lingual root face. The end
of the distal root lobe and the entire mesial root lobe are
missing. The lingual protuberance is well developed and
contains a deep nutrient groove. Multiple small foramina
are situated throughout the preserved portion of the root.
Lateral teeth possess a median cusp that is distally
inclined. The labial crown face is flat and does not
overhang the labial root face. Short and strong enameloid
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Turonian Euselachians from Canada 571
Figure 11. Scapanorhynchus, odontaspidid and Cretalamna teeth from the Watino localities. A, Scapanorhynchus sp., fractured lateral
tooth, UALVP 53193. B–E, Carcharias aff. C. striatula (Dalinkevicius 1935); B, fractured anterior tooth, UALVP 53194; C, lateral tooth,
UALVP 53195; D, lateral tooth, UALVP 53196; E, lateral tooth, UALVP 53197; F–I, Odontaspis watinensis sp. nov.; F, anterior tooth,
UALVP 53199 (holotype); G, fractured anterior tooth, UALVP 53200; H, fractured lateral tooth, UALVP 53201; I, fractured lateral tooth,
UALVP 53202; J–M, Johnlongia parvidens (Cappetta 1973); J, fractured anterior tooth, UALVP 53203; K, fractured lateral tooth, UALVP
53204; L, fractured anterior tooth, UALVP 53205; M, lateral tooth, UALVP 53206. N, Cretalamna ex. gr. appendiculata, fractured lateral
tooth, UALVP 53208. Views: labial (left), profile (centre), and lingual (right) for A, B, F, and N; labial (left), profile (centre), and occlusal
(right) for J; labial (top) and lingual (bottom) for M; labial (left) and lingual (right) for C–E, G–I, K and L. E, F, H–K, and M are SEM
images. Scale bars =1 mm.
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572 T. D. Cook et al.
folding is present along the base of the labial crown. The
lingual crown face is convex and contains very weak
striations that are restricted to the lower half of the cusp.
Flanking the median cusp is a pair of short triangular
lateral cusplets. A distinct cutting edge runs continuously
between the median cusp and lateral cusplets. A narrow
lingual neck is present. The root is relatively robust and
more or less symmetrical. The basal concavity is V-shaped
and is relatively shallow in distally positioned lateral
teeth. The lingual protuberance contains a well-developed
nutrient groove. Numerous large foramina are present in
the medial region of the labial root face. Additional small
foramina are situated throughout the root.
Remarks. Glickman (1980) erected Eostriatolamia to
accommodate odontaspidid-like teeth originally described
as Lamna venusta by Leriche (1906) from the Santonian of
Lonz´
ee, Belgium. Tooth morphology included the presence
of short striations along the base of the labial crown face
and weak striations on the lower half of the lingual crown
face. Glickman & Averianov (1998) noted the importance
of the lingual striations as a principal character distin-
guishing this taxon from other odontaspidids and assigned
additional species (including Carcharias striatula)to
Eostriatolamia based on this character. However, the taxo-
nomic significance of this feature is questionable as lingual
striations also occur in Carcharias (Purdy 1998). Glickman
& Averianov (1998) also stated that despite having similar
tooth morphology to the extant species Carcharias taurus
Rafinesque, 1810, the dentition of Eostriatolamia is more
primitive as it lacks distally positioned files of reduced
crushing teeth. This assertion was based on the failure
to recover the isolated teeth of this morphology during
sampling. It is possible, however, that “physical parameters
acting on the sediment” may separate small distal teeth
from larger more medially positioned teeth (Siverson 1992,
p. 536). Underwood & Cumbaa (2010) also supported
the creation of Eostriatolamia by reassigning Carcharias
tenuiplicatus (Cappetta & Case 1975) and C. striatula
to this genus. In addition, they erected a new species,
E. paucicorrugata Underwood & Cumbaa, 2010. Given
that the validity of Eostriatolamia is based on the absence
of reduced crushing teeth, the assignment of this new
species to Eostriatolamia is questionable as it was noted
that there was a “lack of sufficient material to reconstruct
the dentition” and that the dental arrangement would be
similar to Odontaspis ferox (Risso, 1810) (Underwood &
Cumbaa 2010, p. 918). We caution that in the absence of
an articulated dentition confirming this primitive condition
(i.e. the lack of files of reduced posterior teeth) we consider
Eostriatolamia to be highly provisional. Consequently, we
conservatively assign odontaspid-like dentition herein to
either Carcharias or Odontaspis Agassiz, 1838. According
to Compagno & Follett (1986, pp. 89, 90), the anterior
teeth of the former have median cusps that are “stout and
broad-tipped” and cusplets that are “short and strongly
hooked”, whereas teeth from equivalent files in the latter
have median cusps that are “slender and narrow-tipped”
and cusplets that are “long and straight or weakly curved”.
The lateral teeth of Carcharias are also more “compressed
and blade-like, with flattened cusplets”.
Carcharias striatula was originally described as Odon-
taspis (Synodontaspis)striatula from the late Albian Jiesia
Formation of Lithuania (Dalinkevicius 1935). It has also
been recovered from the late Aptian of southern France
(Cappetta 1975; as Odontaspis striatula), the middle
Albian of northeastern France (Landemaine 1991; as
Scapanorhynchus striatula; Biddle 1993) and the middle
to late Albian of Western Australia (Siverson 1997). The
teeth of this species were reported to be morphologically
similartothoseofC. tenuiplicatus (Welton & Farish 1993);
however, Cappetta & Case (1999) noted that the teeth of
C. striatula are thicker and larger in overall size, have a
straighter median cusp, and less developed striations of the
lingual crown face.
The Watino specimens differ only slightly from the older
Carcharias striatula type material in having anterior teeth
with a more robust and shorter median cusp with less devel-
oped labial striations and lateral teeth with a higher lingual
root face. However, the overall morphology most likely
suggests that the Watino specimens are conspecific. This
material can also be distinguished from C. tenuiplicatus by
having a larger overall size, erect median cusps on anterior
teeth, weak labial and lingual crown striations and shallower
labial basal ledge. Consequently, we consider the Watino
teeth to be congeneric but not conspecific with C. tenuipli-
catus. Teeth identified as Carcharias sp. A from the basal
Haycock Marl of Wester n Australia (Siverson 1996) are also
somewhat similar to the Watino specimens but bear stronger
lingual striations and have a more gracile morphology.
Genus Odontaspis Agassiz, 1838
Odontaspis watinensis sp. nov.
(Fig. 11F–I)
Diagnosis. Teeth of Odontaspis watinensis can be distin-
guished from the teeth of other odontaspidids by possessing
a single pair of needle-like lateral cusplets that extend
nearly half the height of the median cusp. The median cusp
contains no ornamentation. A well-developed and slightly
recessed lingual neck is present on all cusps. The bilobate
root with a well-developed lingual protuberance possesses
a shallow nutrient groove.
Derivation of name. Named after the town of Watino,
Alberta, Canada, near the type locality.
Material. Holotype: UALVP 53199, anterior tooth, local-
ity 76. Paratypes: UALVP 53200, fractured anterior tooth,
locality 76; UALVP 53201, fractured lateral tooth, locality
76; UALVP 53202, fractured lateral tooth, locality 76.
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Turonian Euselachians from Canada 573
Description. Anterior teeth possess an erect median cusp
that tapers to a sharp apex. The labial face is flat, whereas
the lingual face is strongly convex. Both faces are smooth.
The cutting edge is continuous with a pair of extremely tall
needle-like lateral cusplets that extend to half the height of
the median cusp. A well-developed and slightly recessed
lingual neck clearly separates the median cusp and lateral
cusplets from the root lingually. The basal ledge of the labial
crown face is concave and slightly overhangs the labial
root face. The slightly more elongated mesial root lobe
is separated from the distal root lobe by a deep U-shaped
basal concavity. A very shallow nutrient groove is situated
on the basal region of the lingual protuberance. Numerous
large foramina are concentrated throughout the labial root
face.
The lateral teeth have a median cusp that is more robust
than anteriorly positioned teeth and is distally curved.
The labial crown face is flat whereas the lingual face is
convex. No ornamentation is present. A well-developed
and slightly recessed lingual neck is present on all cusps.
The basal ledge is arched and slightly overhangs the labial
root face. The bilobate root forms a deep basal concavity
even in the more distally positioned lateral teeth.
Remarks. The overall morphology of Odontaspis wati-
nensis is somewhat reminiscent of three odontaspidids
occurring in Cenomanian and Turonian deposits of the
WIS: O. saskatchewanensis Case, Tokaryk & Baird,
1990, Carcharias tenuiplicatus and C. paucicorrugata.
The tooth morphology of O. watinensis differs from O.
saskatchewanensis by having: (1) a narrower median cusp;
(2) significantly taller lateral cusplets; and (3) an overall
larger size. Case et al. (1990) gave an average tooth height
of 3.5 mm for the latter species. Welton & Farish (1993)
reported O. saskatchewanensis (as Carcharias sp. A) as
having a maximum tooth size of 4.7 mm, while Cicimurri
(2001a, b) described numerous teeth all measuring less than
5 mm in total height. Shimada et al. (2006) examined more
than 300 specimens of O. saskatchewanensis with no teeth
exceeding 5 mm in total height. The height of O. watinensis
anterior teeth reported herein greatly exceeds the height of
any previously reported O. saskatchewanensis teeth.
The tooth morphology of Odontaspis watinensis differs
from Carcharias tenuiplicatus by possessing: (1) signif-
icantly taller lateral cusplets; (2) completely smooth labial
and lingual crown faces; (3) a labial crown basal ledge that
does not overhang the labial root face to the same degree;
and (4) a larger overall size.
The teeth of Odontaspis watinensis differs from
Carcharias paucicorrugata by having: (1) anterior teeth
with a narrower median cusp; (2) lateral cusplets that are
significantly taller and less divergent; (3) a labial crown
face that is completely smooth; (4) a less developed labial
basal ledge; (5) a single pair of lateral cusplets; and (6) a
shallower nutrient groove.
Odontaspis watinensis tooth morphology bears some
resemblance to O. subulata (Agassiz, 1843, pl. 37a, figs 5,
6; as Lamna subulata) originally described from the
Cenomanian of Germany. Both species have teeth with
smooth crowns and needle-like cusps; however, the latter
has a straight labial crown base, shorter cusplets, and more
elongated root lobes.
Teeth with similar morphology have not been reported
from any other known deposit.
Genus Johnlongia Siverson, 1996
Johnlongia parvidens (Cappetta, 1973)
(Fig. 11J–M)
Material. UALVP 53203, fractured anterior tooth, locality
76. UALVP 53204, fractured lateral tooth, locality 76.
UALVP 53205, fractured anterior tooth, locality 76.
UALVP 53206, lateral tooth, locality 76. UALVP 53207,
10 fractured teeth, locality 848.
Description. Anterior teeth have a tall and slender median
cusp that is slightly distally inclined and strongly lingually
directed. The convex labial face and strongly convex
lingual face are smooth. The cutting edge is restricted
to the apical third of the cusp. The lateral cusplets are
missing but are needle-like in specimens recovered from
other deposits. The lingual neck is narrow. The root
contains a massive lingual protuberance that is bisected
by a deep nutrient groove and has numerous foramina.
The mesial lobe is typically more elongated than the distal
lobe.
Lateral teeth have a broad-based median cusp that tapers
to a sharp apex and is distally inclined. The slightly convex
labial crown face has numerous strong enameloid folds that
are restricted to the crown base. The lingual face is strongly
convex and generally smooth. The mesial lateral cusplet
is tall and needle-like, whereas the distal lateral cusplet is
missing in the figured specimen. The root is asymmetrical
and is missing the distal root lobe and the terminal portion
of the mesial root lobe. The lingual protuberance has
a well-developed nutrient groove and is significantly
more labiolingually compressed compared to anteriorly
positioned teeth. The median cusp and lateral cusplets are
expanded mesodistally in distally positioned lateral teeth.
Remarks. The Watino specimens are the second docu-
mented report of Johnlongia parvidens from Canada. The
first account was a single anterior tooth recovered from the
Dunvegan Formation of Alberta (Cook et al. 2008). The
anterior teeth described herein have a similar morphology
to this tooth.
Siverson (1996) erected Johnlongia allocotodon based
on teeth collected from latest Cenomanian/earliest Turo-
nian deposits of Western Australia. According to Siverson
(1996), the teeth of this species were readily distinguishable
from the late Turonian J. parvidens type material (Cappetta
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574 T. D. Cook et al.
1973) by the anterior teeth having a longer distal cutting
edge, lateral teeth bearing ornamentation on the labial
crown face and the teeth having an overall larger size.
A comparison of the distally positioned lateral teeth of
these species could not be made because teeth from the
posterior half of the jaw of J. parvidens were not included
in the description of the type material (Siverson 1996).
The significance of these characters and the validity of
J. allocotodon were discussed in Cook et al. (2008).
The lateral teeth recovered from the Watino localities
have strong labial crown face ornamentation and a rela-
tively labiolingually compressed root similar to the lateral
teeth of Johnlongia allocotodon. As well, the anterior teeth
described herein also have a distal cutting edge of similar
length and are comparable in overall size to the Australian
material (see Siverson 1996, pl. 5). Although it is possible
that both the Dunvegan and the Watino teeth belong to J.
allocotodon, until re-examination of the type material and
the recovery of additional topotypic teeth can be made, we
conservatively assign these teeth to J. parvidens.
Fam ily Otodontidae Glickman, 1964
Genus Cretalamna Glickman, 1958
Cretalamna ex. gr. appendiculata (Agassiz, 1835)
(Fig. 11N)
Material. UALVP 53208, fractured lateral tooth, locality
76.
Description. The lateral tooth contains a broad triangular
median cusp that is labiolingually compressed and distally
inclined. The labial crown face is slightly convex, whereas
the lingual crown face is more strongly convex. Both faces
are smooth. A distinct cutting edge runs continuously across
the median cusp and a pair of triangular lateral cusplets. The
preserved portion of the distal root lobe appears to be more
robust than the rectangular-shaped mesial root lobe. A shal-
low U-shaped basal concavity separates the lobes. There is a
very weak lingual protuberance that lacks a nutrient groove.
Remarks. The teeth of Cretalamna appendiculata species
group have been reported from various North American
deposits including: the Albian through Maastrichtian of
Texas (Welton & Farish 1993); the Turonian of New
Mexico (Wolberg 1985); the late Cenomanian and Turo-
nian of Arizona (Williamson et al. 1993); the middle
Cenomanian of Colorado (Shimada et al. 2006); the late
Cenomanian of Kansas (Shimada & Martin 2008); and the
late Cenomanian and middle Turonian of South Dakota
and Wyoming (Cappetta 1973; Cicimurri 2001a, 2004).
Cumbaa et al. (2006) reported the teeth of this species from
the Carrot River bone bed of Saskatchewan. Underwood &
Cumbaa (2010, p. 916) noted that these teeth, along with
a single tooth recovered from the Bainbridge bone bed,
differ from C. appendiculata by “possessing a more robust
root with a far more rounded profile”. As such, these teeth
were left in open nomenclature. They also suggest that
teeth allocated to C. appendiculata from different localities
are of variable morphology and most likely represent
multiple species, a view shared herein. Compared to the
Saskatchewan material, UALVP 53208 has an angular root
morphology that is more reminiscent of C. appendiculata.
However, more and better-preserved material will need
to be recovered from the Watino localities before we can
conclusively report the presence of C. appendiculata.
Despite this taxonomic uncertainty, Cretalamna material
from the mid Cretaceous of Canada appears to be rare
(Cumbaa et al. 2006; Underwood & Cumbaa 2010).
Fam ily Incertae familiae
Genus Cretodus Sokolov, 1965
Cretodus semiplicatus (M¨
unster in Agassiz, 1843)
(Fig. 12A)
Material. UALVP 53209, fractured lateral tooth, locality
76; UALVP 53210, fractured lateral tooth, locality 76.
Description. The lateral tooth contains a tall, triangular
median cusp that is slightly distally inclined. The labial
crown face is slightly convex, whereas the lingual crown
face is strongly convex. A pair of divergent triangular
lateral cusplets flanks the median cusp. A distinct cutting
edge runs continuously between the cusps. Strong vertical
enameloid folding is restricted to the basal half of the
median cusp and lateral cusplets labially. The lingual face
of the median cusp and lateral cusplets contains folds that
approach the apex. The distinct lingual neck is narrow.
The root is damaged and is missing a large portion of
the distal lobe and the terminal region of the mesial lobe.
The root lobes are separated by a deep basal concavity.
The well-developed lingual protuberance lacks a nutrient
groove but contains a cluster of large foramina.
Remarks. Cook et al. (2008) reported the recovery
of juvenile teeth of Cretodus semiplicatus from the
middle Cenomanian Dunvegan Formation of Alberta. The
recovery of C. semiplicatus teeth from various middle
Cenomanian–middle Turonian deposits throughout the
WIS was summarized in that paper. Cumbaa et al. (2006,
fig. 4.11) figured posterior teeth of this species from the
Bainbridge bone bed of Saskatchewan. Subsequently,
Underwood & Cumbaa (2010) suggested that these
posterior teeth in fact belong to Archaeolamna ex. gr.
kopingensis, an observation shared herein.
Teeth from the congeneric Cretodus longiplicatus were
first described by Werner (1989) from the late Cenomanian
of Bahariya, Egypt. Later, Cappetta & Case (1999)
synonymised this taxon with C. semiplicatus. Based on
teeth recovered from the early Cenomanian of India,
Underwood et al. (2011, p. 548) reinstated C. longiplicatus
as a valid species based on the “long and very robust
longitudinal ridges on both labial and lingual faces of the
crown” present throughout the ontogeny of the species.
They proposed that this crown ornamentation was less
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Turonian Euselachians from Canada 575
Figure 12. Cretodus,Dallasiella and Rhinobatos teeth from the Watino localities. A, Cretodus semiplicatus (M¨
unster in Agassiz 1843),
fractured lateral tooth, UALVP 53209; B–H, Dallasiella willistoni Cappetta & Case, 1999; B, lateral tooth, UALVP 53211; C, fractured
lateral tooth, UALVP 53212; D, fractured anterior tooth, UALVP 53213; E, fractured lateral tooth, UALVP 53214; F, fractured lateral
tooth, UALVP 53215; G, lateral tooth, UALVP 53216; H, fractured lateral tooth, UALVP 53217; I, J, Rhinobatos incertus Cappetta, 1973;
I, tooth from female, UALVP 53220; J, tooth from male, UALVP 53221. Views: labial (left), profile (centre), and lingual (right) for A and
C; labial (top) and lingual (bottom) for G and H; occlusal (left) and labial (right) for I; occlusal (left), and profile (right) for J; labial (left)
and lingual (right) for B, and D–F. B, I and J are SEM images. Scale bars for A–H =1 mm; scale bars for I and J =0.5 mm.
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576 T. D. Cook et al.
developed and maximum tooth size was larger in the
younger C. semiplicatus. Underwood et al. (2011, p. 549)
also stated that C. longiplicatus was “absent north of Tethys
in either Eurasia or the Western Interior Seaway”. The
partially complete C. longiplicatus anterior tooth figured
by Underwood et al. (2011, fig. 8e, f) is only slightly
larger in size than the C. semiplicatus specimen described
herein. Both the Watino and Indian teeth have a similar
overall morphology and pattern of enameloid folding. A
juvenile tooth from the middle Cenomanian of Alberta
also has prominent enameloid folds that extend most of
the length of the crown. Consequently, the revalidation of
C. longiplicatus based on the length of crown folding and
stratigraphy is contradicted by the Alberta specimens and
this taxon is considered invalid herein. Differences in the
length of enameloid folding between early–middle Turo-
nian occurrences of C. semiplicatus is likely attributable to
intraspecific variation (Underwood et al. 2011).
Genus Dallasiella Cappetta & Case, 1999
Dallasiella willistoni Cappetta & Case, 1999
(Fig. 12B–H)
Material. UALVP 53211, lateral tooth, locality 76.
UALVP 53212, fractured lateral tooth, locality 848. UALVP
53213, fractured anterior tooth, locality 848. UALVP
53214, fractured lateral tooth, locality 848. UALVP 53215,
fractured lateral tooth, locality 76. UALVP 53216, lateral
tooth, locality 848. UALVP 53217, fractured lateral tooth,
locality 76. UALVP 53218, 10 fractured teeth, locality 848.
Description. Anterior teeth have a tall narrow median cusp
that is more or less erect and is slightly lingually directed.
The cusp has a more or less flat labial face and a strongly
convex lingual face. Both faces are smooth. The cutting
edge runs continuously between the median cusp and a pair
of tall triangular lateral cusplets. The height of the narrow
lingual neck is constant between the median cusp and lateral
cusplets. The sigmoidal basal ledge of the labial crown
face overhangs the labial root face. The root is more or
less symmetrical and contains root lobes that are distinctly
labiolingually compressed. The lingual protuberance is
very well developed and bears a large nutrient foramen that
is situated in a small nutrient groove. Additional foramina
are present throughout the labial root face, particularly
concentrated in the medial region below the crown root
border. The basal concavity is deep and U-shaped.
Lateral teeth have a narrow median cusp that is distally
inclined. The labial crown face is more or less flat, whereas
the lingual crown face is strongly convex. Both faces are
smooth. A distinct cutting edge runs continuously across the
median cusp and a pair of broad triangular lateral cusplets.
A second mesial cusplet may be present in distally posi-
tioned lateral teeth. The lingual neck is narrow. The basal
ledge only slightly overhangs the labial root face and is
sigmoidal. The mesial and distal root lobes are more or
less symmetrical, divergent, and labiolingually compressed.
The lingual protuberance is well developed and contains a
deep nutrient groove that houses a large nutrient foramen.
Numerous smaller foramina are concentrated below the
crown root border on the labial root face. The basal concav-
ity is shallow in the most distally positioned lateral teeth.
Remarks. Dallasiella willistoni has been reported from
the Turonian–Coniacian of Texas (Cappetta & Case 1999)
and the middle Turonian of Montana (Siverson & Lindgren
2005). The general tooth morphology of Dallasiella is
similar to that of Cretalamna;however,the latter lacks a
deep nutrient groove (Cappetta & Case 1999). The labiolin-
gual compressed root lobes also differentiate this species
from juvenile Archaeolamna teeth, which may also possess
a nutrient groove, although less developed (Underwood &
Cumbaa 2010). The robust root morphology suggests that
the tooth reported as D. willistoni by Cook et al. (2008)
is more likely Archaeolamna ex. gr. kopingensis. A tooth
described as Cretodus sp. by Case et al. (1990, fig. 7e, f)
from the lower Turonian Favel Formation of Saskatchewan
strongly resembles the Watino specimens and is considered
conspecific herein.
Order Rajiformes Berg, 1940
Fam ily Rhinobatidae M¨
uller & Henle, 1838
Genus Rhinobatos Linck, 1790
Rhinobatos incertus Cappetta, 1973
(Fig. 12I, J)
Material. UALVP 53220, female tooth, locality 76.
UALVP 53221, male tooth, locality 76. UALVP 53222, 79
teeth, locality 76. UALVP 53223, 109 teeth, locality 848.
Description. Teeth are extremely small. The crown is
mesodistally elongated and lacks ornamentation. The labial
face is slightly convex and contains a rounded crown foot.
A distinct lingually directed narrow cusp may be present
(UALVP 53221) or may be reduced or absent (UALVP
53220). In the non-cuspate condition, there is a cutting
edge separating the labial and lingual crown faces. In the
cuspate condition, this cutting edge is not defined. The
lingual crown face bears a narrow and elongated central
uvula which is separated from less developed lateral uvulae
by deep grooves. The root is displaced slightly lingually
below the crown and is divided into two lobes by a deep
nutritive groove containing a large foramen. Additional
foramina are located on the basal surface of each root lobe
and on the root face on either side of the central uvula.
Remarks. TheteethofRhinobatos are sexually dimor-
phic. Males in the breeding phase possess a crown with
a distinct sharp and elongated cusp, whereas females and
non-breeding males lack this structure (Kajiura & Tricas
1996; Everhart 2007). Cappetta & Case (1999) noted that
R. incertus can be differentiated from other species of
Rhinobatos by its distinct cusp, lack of sharp cutting edges,
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Turonian Euselachians from Canada 577
Tab l e 1 . List of euselachian taxa in the early Turonian Watino assemblage described herein, and comparisons with the early Turonian
Favel assemblage (Case et al. 1990) and middle Cenomanian Bainbridge bonebed assemblage (Underwood & Cumbaa 2010) of
Saskatchewan, and the middle Cenomanian Dunvegan Formation assemblage (Cook et al. 2008) of Alberta, Canada.
Early Turonian Middle Cenomanian
Alberta Saskatchewan Alberta Saskatchewan
(Watino) (Favel) (Dunvegan) (Bainbridge)
Meristodonoides rajkovichiaX– X X
Polyacrodus sp. X – – –
Ptychodus anonymusbXX – –
Ptychodus ex. gr. decurrens –– – X
Ptychodus rhombodus –– – X
Ptychodus sp. – – – X
Squalicorax sp. AcXXXX
Squalicorax sp. B X – – –
Palaeoanacorax aff. pawpawensis –– – X
Archaeolamna ex. gr. kopingensisdX– X X
Cardabiodon aff. C. ricki X– – –
Cretoxyrhina mantellieXXXX
Protolamna carteri –– X –
Scapanorhynchus sp. X – – –
Carcharias lilliae –X – –
Carcharias paucicorrugataf–– – X
Carcharias aff. C. striatula X– – –
Odontaspis saskatchewanensis –X – –
Odontaspis watinensis nov. sp. X – – –
Johnlongia parvidens X– X –
Roulletia canadensis –– – X
Cretalamna ex. gr. appendiculatagX– – X
Cretodus semiplicatus X– X –
Dallasiella willistonihXX – –
Orectoloboides angulatus –– – X
Cretorectolobus robustus –– – X
Pseudohypolophus mcnultyi –– X –
Rhinobatos incertusiXX – X
Cretomanta canadensis –X – X
aReported as Hybodus sp. by Cook et al. (2008); incomplete teeth reported herein as Meristodonoides cf. rajkovichi.bReported as Ptychodus cf. P. rugosus
by Case et al. (1990). cReported as Squalicorax falcatus by Case et al. (1990); reported as S. curvatus by Underwood & Cumbaa (2010); reported as
Squalicorax sp. by Cook et al. (2008). dReported as Dallasiella willistoni by Cook et al. (2008). eTeeth reported as Cretodus sp. by Case et al. (1990, fig.
7a–d) are likely juvenile Cretoxyrhina mantelli; reported as Cretoxyrhina cf. C. mantelli by Cook et al. (2008); Underwood & Cumbaa (2010) reported
teeth of similar morphology but containing lateral cusplets in anteriorly positioned teeth as C. denticulata.fReported as Eostriatolamia paucicorrugata
by Underwood & Cumbaa (2010). gReported as Cretalamna sp. by Underwood & Cumbaa (2010). hReported as Cretodus sp. by Case et al. (1990, fig.
7e, f). iReported as Rhinobatos sp. by Case et al. (1990); reported as ‘Rhinobatos’cf.incertus by Underwood & Cumbaa (2010).
and lateral uvulae that are weakly developed. These charac-
teristics are observed in the male specimens recovered from
Watino.
Rhinobatos incertus has been described from North
American localities including the lower Turonian and
Coniacian of Texas (Welton & Farish 1993), lower Turo-
nian of Arizona (Williamson et al. 1993; as Rhinobatos
sp.), middle Cenomanian of Colorado (Shimada et al.
2006; as Rhinobatos sp.), late Albian to early Campanian
of Kansas (Shimada & Martin 2008; Everhart 2007),
late Cenomanian and middle and late Turonian of South
Dakota (Cappetta 1973; Cicimurri 2001a, 2004), and
middle Cenomanian (Underwood & Cumbaa 2010; as
Rhinobatos cf. incertus) and lower Turonian (Case et al.
1990; as Rhinobatos sp.) of Saskatchewan.
Discussion
The assemblage described herein includes 16 species
belonging to at least three orders and 11 families and
15 genera, with the most abundant remains belonging to
two different species of Squalicorax. The first description
is given of the new species Odontaspis watinensis,
and the first known occurrences of Polyacrodus sp.,
Scapanorhynchus sp. and Carcharias aff. C.striatula from
Canada are recorded, extending the northern geographical
ranges of these taxa within the WIS.
The Watino euselachian assemblage has species in
common with an early Turonian assemblage recovered
from the Keld Member of the Favel Formation in east-
central Saskatchewan (Case et al. 1990) (Table 1). The latter
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578 T. D. Cook et al.
assemblage was situated at a relatively similar palaeolati-
tude and deposited in an offshore environment (Cumbaa
& Tokaryk 1999; Cumbaa et al. 2006; Wu et al. 2001).
Both the Watino and Favel assemblages share at least three
taxa (Ptychodus anonymus,Squalicorax sp. A and Rhino-
batos incertus). As mentioned above, a tooth identified
as Cretodus sp. by Case et al. (1990, fig. 7e, f) strongly
resembles Dallasiella willistoni. Two additional Cretodus
sp. teeth (Case et al. 1990, fig. 7a–d) likely belong to a small
juvenile Cretoxyrhina mantelli. The Saskatchewan assem-
blage has two species (Carcharias lilliae (Case, Tokaryk &
Baird, 1990) and Cretomanta canadensis Case, Tokaryk &
Baird, 1990)) that have not been recovered from Watino.
Conversely, 10 species (Meristodonoides rajkovichi, Poly-
acrodus sp., Squalicorax sp. B, Cardabiodon aff. C. ricki,
Scapanorhynchus sp., Carcharias aff. C. striatula,Odon-
taspis watinensis,Johnlongia parvidens,Cretalamna ex. gr.
appendiculata, and Cretodus semiplicatus) are present in
the Watino assemblage but absent in the Favel assemblage.
Temperature and salinity are the principal physi-
cal factors influencing extant selachian distributions
(Simpfendorfer & Heupel 2004). Accordingly, habitat
selection may have contributed to the difference in
the faunal composition between the Watino and Favel
assemblages. Despite only a 6◦separation in palaeolatitude
(PLATES project 2010), sea surface temperature along
the two coasts of the WIS probably differed. Using model
simulations, Slingerland et al. (1996) proposed a counter-
clockwise estuarine circulation for the WIS. The outflow
of freshwater drainage drew cooler Boreal water down
the western margin and carried warmer Tethyan water up
the eastern shoreline, resulting in distorted isotherms. It is
possible that there may have been a 6◦C difference in water
temperature between the Watino and Favel localities during
the early Turonian (see Slingerland et al. 1996, fig. 6).
The northward movement of warm Tethyan water along
the eastern side of the WIS is supported by foraminiferal
distribution studies (Eicher & Diner 1985; Fisher et al.
1994; Schr¨
oder-Adams et al. 1996, 2001). Temperature
intolerance to warmer water may account for the absence
of Carcharias aff. C. striatula,Odontaspis watinensis, and
Cardabiodon aff. C. ricki from the Favel assemblage, as
these species have not been recovered from additional WIS
assemblages situated in presumed warmer WIS isotherms.
Conversely, the absence of Odontaspis saskatchewanensis,
Carcharias lilliae, and Cretomanta canadensis from
Watino may suggest that these species avoided cooler
waters. It should be noted that Odontaspis watinensis and
Carcharias lilliae have not been reported outside of the
Watino and Favel assemblages, respectively, which may
indicate that these species were endemic to these particular
regions. Temperature intolerance, however, cannot explain
the absence of Meristodonoides rajkovichi,Polyacrodus
sp., Squalicorax sp. B, Scapanorhynchus sp., Johnlongia
parvidens,Cretalamna ex. gr. appendiculata, and Cretodus
semiplicatus from the Favel assemblage, as these taxa
have been recovered from contemporaneous assemblages
in similar or warmer isotherms of the WIS. Salinity, an
additional factor shaping elasmobranch distribution, does
not appear to be markedly different between the two
localities according to the model (see Slingerland et al.
1996, fig. 8) and likely did not contribute to the variation in
diversity.
Biotic factors, such as the availability of prey, also
influence elasmobranch distributions (Heithaus et al.
2002; Simpfendorfer & Heupel 2004). Associated with
the Favel euselachian assemblage, Cumbaa & Tokaryk
(1999) reported remains of the teleosts Xiphactinus audax
Leidy, 1870, Ichthyodectes,Gillicus Hay, 1898, Apsopelix
anglicus (Dixon, 1850), Pachyrhizodus minimus Stewart,
1899, Enchodus and Protosphyraena Leidy, 1857b. From
the Watino sandstone, Wilson & Chalifa (1989) reported
Belonostomus cf. B. longirostris (Lambe, 1902), Ichthyo-
dectes ctenodon Cope, 1870, Apateodus sp., Cimolichthys
cf. C. levesiensis Leidy, 1857a, cf. Dercetoides sp. and
Enchodus cf. E. shumardi Leidy, 1857a. Additional early
Turonian taxa were recovered from Watino concretions and
included cf. Gillicus sp., cf. Xiphactinus sp., Osmeroides
cf. O. delicatus (Cockerell, 1919) and Leucichthyops
sp. Both the Watino and Favel localities have numerous
teleosts, as juvenile and/or adult forms, that could serve
as potential prey items. Therefore, a disparate food source
fails to adequately explain the variation in euselachian
diversity between the Watino and Favel assemblages.
With the exception of Squalicorax sp. B., the afore-
mentioned taxa absent from the Favel assemblage are only
represented by rare elements in the Watino euselachian
assemblage. Hence, the difference in faunal composition
may be the result of collecting bias. The absence of Squal-
icorax sp. B teeth in the Favel assemblage is puzzling, as
it is among the most abundant euselachian material recov-
ered from Watino and has been reported from the middle
Cenomanian of Saskatchewan (Cumbaa et al. 2006; as
S. volgensis).
Comparing the early Turonian Watino assemblage with
the middle Cenomanian assemblages recovered from
the Bainbridge bone bed of east-central Saskatchewan
(Underwood & Cumbaa 2010) and the Dunvegan Forma-
tion of northwestern Alberta (Cook et al. 2008), we are
able to document changes in faunal composition from
middle Cenomanian to early Turonian in the northern
region of the seaway (Table 1). The Bainbridge assemblage,
recovered from the Bell Fourche Member of the Ashville
Formation, contains five species (Meristodonoides
rajkovichi,Squalicorax sp. A, Archaeolamna ex. gr.
kopingensis,Cretalamna ex. gr. appendiculata, and
Rhinobatos incertus) in common with the Watino
assemblage. Eight species (Ptychodus ex. gr. decurrens
Agassiz, 1838, Ptychodus rhombodus,Palaeoanacorax
aff. pawpawensis,Carcharias paucicorrugata,Roulletia
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Turonian Euselachians from Canada 579
canadensis Underwood & Cumbaa, 2010, Orectoloboides
angulatus Underwood & Cumbaa, 2010, Cretorectolobus
robustus Underwood & Cumbaa, 2010, and Cretomanta
canadensis) are present in the Bainbridge assemblage
but absent in the Watino assemblage. Conversely, eight
species (Polyacrodus sp., Squalicorax sp. B, Cardabiodon
aff. C. ricki,Scapanorhynchus sp., Carcharias aff. C.
striatula,Odontaspis watinensis,Cretodus semiplicatus,
and Dallasiella willistoni) are present in the Watino
assemblage but absent in the Bainbridge assemblage.
Both assemblages also contain Cretoxyrhina; however, the
anterior teeth from Watino lack lateral cusplets.
The Dunvegan assemblage is believed to represent a
marginal marine fauna that likely endured periodic fluctua-
tions in salinity due to the influx of fresh water (Cook et al.
2008). Although the Dunvegan assemblage is considerably
less productive than the Watino assemblage, five species
(Meristodonoides rajkovichi,Squalicorax sp. A, Archaeo-
lamna ex. gr. kopingensis,Cretodus semiplicatus and John-
longia parvidens) are common to both assemblages. Two
species (Protolamna carteri and Pseudohypolophus mcnul-
tyi) are present in the Dunvegan assemblage but absent in
the Watino assemblage, whereas 10 species (Polyacrodus
sp., Ptychodus anonymus,Squalicorax sp. B, Cardabiodon
aff. C. ricki,Scapanorhynchus sp., Carcharias aff. C.striat-
ula,Odontaspis watinensis,Cretalamna sp., Dallasiella
willistoni, and Rhinobatos incertus) are exclusive to the
latter. Both assemblages also contain Cretoxyrhina.
Of the ten species (Ptychodus ex. gr. decurrens,
Ptychodus rhombodus,Palaeoanacorax aff. P. pawpawen-
sis,Protolamna carteri, Carcharias paucicorrugata,
Roulletia canadensis,Orectoloboides angulatus,Cretorec-
tolobus robustus,Pseudohypolophus mcnultyi, and
Cretomanta canadensis) present in the middle Ceno-
manian Dunvegan and Bainbridge assemblages but
absent in the early Turonian Watino assemblage, five
species (Ptychodus rhombodus,Palaeoanacorax aff.
P. pawpawensis,Protolamna carteri,Orectoloboides
angulatus and Cretorectolobus robustus) have not been
recovered from WIS deposits younger than the middle
Cenomanian. Teeth recovered from early late Cenomanian
deposits of South Dakota (Cicimurri 2001a, fig. 7m)
and Kansas (Shimada & Martin 2008, fig. 5g) and
identified as Carcharias aff. amonensis and Carcharias
amonensis, respectively, have a morphology more similar
to that of Roulletia canadensis. This latter species was
reported to be “extremely common” in the Bainbridge
assemblage (Underwood & Cumbaa 2010, p. 921)
but is absent at Watino. As such, this species may have
been restricted to middle to early late Cenomanian deposits.
Of the seven species (Polyacrodus sp., Squalicorax
sp. B, Cardabiodon aff.C.ricki,Scapanorhynchus sp.,
Carcharias aff. C. striatula,Odontaspis watinensis, and
Dallasiella willistoni) recovered from the early Turonian
Watino assemblage but absent from the middle Cenoma-
nian Dunvegan and Bainbridge assemblages, only two
species (Odontaspis watinensis and Dallasiella willistoni)
have not been reported from other deposits older than the
Cenomanian–Turonian boundary.
In summary, euselachians that occurred before
the Cenomanian–Turonian boundary in the Cana-
dian region of the WIS include: Ptychodus ex. gr.
decurrens,Ptychodus rhombodus,Palaeoanacorax aff.
P. pawpawensis,Carcharias paucicorrugata,Roulletia
canadensis,Protolamna carteri,Orectoloboides angulatus,
Cretorectolobus robustus and Pseudohypolophus mcnul-
tyi. Euselachians present after the Cenomanian–Turonian
boundary in this region include: Polyacrodus sp., Ptychodus
anonymus,Squalicorax sp. B, Cardabiodon aff. C.
ricki,Scapanorhynchus sp., Carcharias aff. C. striat-
ula,Odontaspis watinensis, and Dallasiella willistoni.
Taxa with a biostratigraphical range extending across the
Cenomanian–Turonian boundary in the northern region
include: Meristodonoides rajkovichi,Squalicorax sp. A,
Archaeolamna ex. gr. kopingensis,Cretoxyrhina,Creta-
lamna ex. gr. appendiculata,Johnlongia parvidens and
Rhinobatos incertus. Including the species recovered
from the aforementioned early Turonian Favel assem-
blage (Case et al. 1990), Cretomanta canadensis spans
the Cenomanian–Turonian boundary, whereas Odontaspis
saskatchewanensis and Carcharias lilliae are reported only
from the early Turonian.
Benthic and/or nektobenthic euselachians are a rare
faunal component in the Watino assemblage. Only two
taxa, Ptychodus and Rhinobatos, reported from this
assemblage likely had a benthic and/or nektobenthic
habitat preference. The former has a grinding-type
tooth morphology indicative of a durophagous lifestyle
(Cappetta 1987); this lifestyle is also supported by the
observation of Shimada et al. (2009, p. 334), who noted
that the scale morphology of Ptychodus occidentalis Leidy,
1868 suggested this species was a “sluggish swimmer that
likely cruised near or at the ocean floor at low speeds.”
The crushing-type dentition of the batoid Rhinobatos
also indicates a benthic habitat (Cappetta 1987). Modern
representatives of this taxon occur in the tropical to
temperate waters of continental shelves (Nelson 2006).
The scarcity of benthic euselachians at Watino and in
other northern assemblages may be due to the persistent
anoxic bottom waters of a stratified water column (Cumbaa
et al. 2010). This stratification may have been the result
of a freshwater layer sitting on top of the marine waters
or the production of a denser intermediate body of water
through the mixing of boreal and Tethyan water masses.
The migration of a planktonic Tethyan fauna into the
northern region of the seaway during the time of peak
transgression resulted in an accumulation of organic
carbon in the bottom waters, which in turn led to an anoxic
environment (Hay et al. 1993; Schr¨
oder-Adams et al. 1996,
2001). Schr¨
oder-Adams et al. (1996) noted that during the
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580 T. D. Cook et al.
Cenomanian–Turonian sea-level high stand, the diversity
of calcareous and agglutinated benthic foraminifera and
nannofossils in the northern region of the WIS was
significantly reduced compared to the southern region of
the WIS, probably because of strong circulation between
the seaway and the Tethys. The rarity of benthic and/or
nektobenthic euselachian taxa at Watino is consistent with
the presence of an anoxic bottom environment. However,
the occurrence of Ptychodus and Rhinobatos indicates
at least some periodic mixing of deeper anoxic waters
with oxygenated surface layers in this region, possibly as
major storms tracked across the seaway allowing benthic
organisms to survive in the area at least occasionally.
Underwood & Cumbaa (2010, p. 940) also noted that
the Bainbridge assemblage was “dominated by pelagic
predators” with few benthic or nektobenthic species. In
addition to Ptychodus and Rhinobatos, they reported the
orectolobiforms Orectoloboides angulatus and Cretorec-
tolobus robustus. The clutching-type tooth morphology of
these species is also consistent with a benthic or nektoben-
thic preference (Case 1978; Cappetta 1987; Bourdon &
Everhart 2010). With the notable exception of the tropical
and warm temperate pelagic species Rhincodon typus
Smith, 1828, extant orectolobiforms are typically tropical,
coastal and benthic (Musick et al. 2004). The rare recovery
of the teeth of benthic species in the Bainbridge fauna was
attributed to infrequent transient individuals to the region
(Underwood & Cumbaa 2010).
Watino shares many species with two lower middle
Turonian assemblages recovered from the Fairport Chalk
Member of the Carlile Formation in western Russell and
southern Ellis counties in north-central Kansas (Figs 13,
14; Table 2). Approximately 3 m of basal Fairport Chalk
and the entire underlying Pfeifer Member of the Greenhorn
Limestone is exposed at the Russell County site. The teeth
from this locality were surface collected from the lower-
most 2 m of the Fairport Chalk in 2007 and 2008 (by MJE).
Along with the euselachian teeth were the remains of Inoce-
ramus cuvieri Sowerby, 1822 and Pseudoperna bentonensis
(Logan, 1899), and small moulds of Collignoniceras wooll-
gari (Mantell, 1822) (MJE pers. obs.). The stratigraphy of
the Ellis County site was described by Everhart & Darnell
(2004) during the description of a Ptychodus mammillaris
tooth recovered from this locality. Fragmentary valves of
Inoceramus cuvieri and Pseudoperna congesta (Conrad
in Nicollet, 1843) were also observed in this exposure.
The Kansas localities are believed to represent an offshore
environment of normal salinity at a probable depth of less
than 100 m (Hattin 1962).
The Kansas teeth figured herein (Figs 13, 14) are
catalogued in the collections at the Fort Hays State
University Sternberg Museum of Natural History (FHSM).
Four species (Squalicorax sp. A, Squalicorax sp. B,
Cretoxyrhina mantelli and Dallasiella willistoni) recovered
from these Kansas localities are also present at Watino.
Only one species (Ptychodus mammillaris) has not been
recovered at Watino. The Cardabiodon material recovered
from Watino belongs to C. aff. C. ricki, whereas the small
juvenile Cardabiodon tooth recovered from Kansas belongs
to the younger congeneric species C. venator. A single
Johnlongia parvidens (FHSM VP-15721) tooth was also
recovered from the Ellis County locality (MJE pers. obs.).
Williamson et al. (1993) reported numerous euselachian
taxa from late Cenomanian–middle Turonian deposits at
Black Mesa, Arizona (Table 2). The early Turonian lower
shale member of the Mancos Shale was deposited below
storm wave-base (Kirkland 1991). Six species (Squal-
icorax sp. A, Cardabiodon sp. (as Cretodus woodwardi,
see Siverson & Lindgren 2005), Cretoxyrhina mantelli,
Scapanorhynchus sp., Cretalamna ex. gr. appendiculata
and Rhinobatos incertus) reported from the lower Turonian
Mancos Shale are also present in the Watino assemblage.
Five species (Ptychodus decurrens,P. marginalis Agassiz,
1839 (as P.cf.P. mammillaris, see Hamm 2010b), P. w h i p -
plei Marcou, 1858, Chiloscyllium greeni (Cappetta, 1973)
and Texatrygon rubyae (Williamson, Kirkland & Lucas,
1993) (as Ptychotrygon rubyae Williamson, Kirkland &
Lucas, 1993, see Cappetta 2006)) are reported from the
Arizona assemblage but are absent at Watino. Conversely,
nine species (Meristodonoides rajkovichi,Polyacrodus sp.,
Ptychodus anonymus,Squalicorax sp. B, Archaeolamna cf.
kopingensis,Carcharias aff. C. striatula,Odontaspis wati-
nensis,Johnlongia parvidens, and Dallasiella willistoni)
are present in the Watino assemblage but absent in the lower
Turonian of Arizona. However, Meristodonoides rajkovichi
(as Hybodus sp., see Underwood & Cumbaa 2010) was
reported from late Cenomanian and middle Turonian
deposits of this southern region (Williamson et al. 1993).
As discussed above, the three Ptychodus species recovered
from Arizona likely had a benthic and/or nektobenthic
habitat. The clutching-type dentition of the orectolobiform
Chiloscyllium greeni and the grinding-type dentition of
the batoid Texatrygon rubyae also suggest a preference
for a benthic habitat (Cappetta 1987). The greater number
of benthic species in this assemblage suggests the bottom
waters of this region were likely more productive, and
oxygenated compared to regions further north.
In summary, seven species (Squalicorax sp. A,
Cretoxyrhina mantelli,Scapanorhynchus sp., Cretalamna
ex. gr. appendiculata,Cretodus semiplicatus,Rhinobatos
incertus and Cretomanta canadensis) have a distribution
within the WIS that spans approximately 20◦of palaeolati-
tude (PLATES project 2010), from Arizona to Alberta and
Saskatchewan, during the early Turonian. All seven species
have also been recovered from Cenomanian and/or Turo-
nian deposits of Texas (Welton & Farish 1993; Cappetta &
Case 1999). Of the additional species recovered from the
Canadian region of the WIS, Meristodonoides rajkovichi,
Polyacrodus sp., Ptychodus anonymus,Squalicorax sp. B
and Odontaspis saskatchewanensis have also been reported
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Turonian Euselachians from Canada 581
Figure 13. Ptychodus and Squalicorax teeth from the lower middle Turonian of Kansas. A, B, Ptychodus mammillaris Agassiz, 1835;
A, medial tooth, FHSM VP-17682; B, lateral tooth, FHSM VP-17683; C–G, Squalicorax sp. A; C, anterior tooth, FHSM VP-17684; D,
anterior tooth, FHSM VP-17685; E, anterior tooth, FHSM VP-17686; F, lateral tooth, FHSM VP-17687; G, lateral tooth, FHSM VP-
17688; H–K, Squalicorax sp. B; H, lateral tooth, FHSM VP-17689; I, lateral tooth, FHSM VP-17696; J, lateral tooth, FHSM VP-17690;
K, lateral tooth, FHSM VP-17697. Views: occlusal (left) and profile (right) for A and B; labial (top) and lingual (bottom) from C; labial
(left) and lingual (right) for D–K. Scale bars =1 mm.
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582 T. D. Cook et al.
Figure 14. Cardabiodon,Cretoxyrhina and Dallasiella teeth from the lower middle Turonian of Kansas. A, Cardabiodon venator Siverson
& Lindgren, 2005, juvenile lateral tooth, FHSM VP-17692; B–F, Cretoxyrhina mantelli (Agassiz, 1843); B, small juvenile lateral tooth,
FHSM VP-17693; C, lateral tooth, FHSM VP-17698; D, lateral tooth FHSM VP-17699; E, lateral tooth, FHSM VP-17700; F, lateral
tooth, FHSM VP-17694; G–I, Dallasiella willistoni Cappetta and Case, 1999; G, anterior tooth, FHSM VP-17701; H, lateral tooth, FHSM
VP-17691; I, lateral tooth, FHSM VP-17695. Views: labial (left) and lingual (right) for A–C and E–I; labial (top) and lingual (bottom)
for D. Scale bars =1 mm.
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Turonian Euselachians from Canada 583
Tab l e 2 . A comparison of the euselachians recovered from the early Turonian of Canada (Case et al. 1990; herein), the lower middle
Turonian of Kansas (herein), and the early Turonian of Arizona (Williamson et al. 1993), USA.
Canada Kansas Arizona
(Watino and Favel) (Fairport Chalk) (Mancos Shale)
Meristodonoides rajkovichiaX––
Polyacrodus sp. X – –
Ptychodus anonymusbX––
Ptychodus decurrens ––X
Ptychodus mammillaris –X–
Ptychodus marginalisc––X
Ptychodus whipplei ––X
Squalicorax sp. AdXXX
Squalicorax sp. B X X –
Archaeolamna ex. gr. kopingensis X––
Cardabiodon aff. C. ricki X––
Cardabiodon venatore–XX
Cretoxyrhina mantelli XXX
Scapanorhynchus sp.fX–X
Carcharias lilliae X––
Carcharias aff. C. striatula X––
Odontaspis saskatchewanensis X––
Odontaspis watinensis nov. sp. X – –
Johnlongia parvidens XX–
Cretalamna ex. gr. appendiculatagXXX
Cretodus semiplicatus X––
Dallasiella willistonihXX–
Chiloscyllium greeni ––X
Rhinobatos incertusiX–X
Texatrygon rubyaej–X
Cretomanta canadensis X––
aIncomplete teeth reported herein as Meristodonoides cf. rajkovichi.; bReported as Ptychodus cf. P. rugosus by Case et al. (1990). cReported as Ptychodus
cf. P. mammillaris by Williamson et al. (1993). dReported as Squalicorax falcatus by Case et al. (1990) and Williamson et al. (1993). eReported as
Cretolamna woodwardi by Williamson et al. (1993). fReported as Scapanorhynchus raphiodon by Williamson et al. (1993). gReported as Cretolamna
appendiculata by Williamson et al. (1993). hReported as Cretodus sp. by Case et al. (1990, fig. 7e, f). iReported as Rhinobatos sp. by Case et al. (1990)
and Williamson et al. (1993). jReported as Ptychotrygon rubyae by Williamson et al. (1993).
from Cenomanian and/or Turonian deposits of Texas
(Welton & Farish 1993; Cappetta & Case 1999). Dallasiella
has also been reported from the Turonian–Coniacian of
Texas (Cappetta & Case 1999). Conversely, Archaeolamna
ex. gr. kopingensis,Cardabiodon aff. C. ricki,Carcharias
aff. C. striatula,Odontaspis watinensis, and Johnlongia
parvidens, have not been documented in the rich Texan
deposits and may have been restricted to cooler waters.
Acknowledgements
We thank M. Siverson (Western Australian Museum),
an anonymous reviewer and the editors for thoughtful
reviews that significantly improved the paper. In addition to
providing stimulating discussions on euselachian systemat-
ics, M. Siverson also identified the Dallasiella willistoni and
small juvenile Cardabiodon teeth from Kansas. The authors
are grateful to A. Lindoe for specimen preparation; D. H.
McNeil (Geological Survey of Canada) for performing a
micropalaeontological analysis on the Watino locality; B.
Varban (Exxon Mobil Upstream Research Geologic Model-
ing, Houston, Texas) and J. I. Kirkland (Utah Geological
Survey) for discussions involving the geology at Watino and
Black Mesa, Arizona, respectively; I. Dalziel and L. Gaha-
gan for providing palaeolatitude data (PLATES Project,
2010, University of Texas Institute for Geophysics, Austin,
Texas); and M. Templin and T. Nomokonova (University of
Alberta) for French and Russian translations, respectively.
We also thank Keith Ewell for assistance in collection of
the Ellis County, Kansas, specimens. This research was
supported by Natural Sciences and Engineering Research
Council of Canada Discovery Grants A9180 to MVHW and
327448 to AMM. Funding and travel grants to MGN were
also provided by the Royal Tyrrell Museum Cooperating
Society.
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