Paludicola 12(2):68-82 June 2019

© by the Rochester Institute of Vertebrate Paleontology

VERTEBRATE TRACE FOSSILS IN THE MOWRY SHALE (LOWER CRETACEOUS)

OF WYOMING, USA

Melissa V. Connely

Department of Earth and Environmental Science, Casper College, Casper Wyoming

mconnely@caspercollege.edu

ABSTRACT
The upper contact of the Mowry Shale Formation (Cretaceous) contains a variety of trace fossils in several locations
throughout Wyoming. Over the past several years, new discoveries of trace fossils have been investigated by students and
faculty at Casper College, Wyoming. Some of the trace fossils discovered are those of vertebrate animals. These trace fossils
resemble swimming and resting marks including claw marks, foot prints, and tail and/or fin drags. These very unique and
poorly known trace patterns may have been produced by marine fish and/or reptiles. At the same level, some impressions have
been identified as the invertebrate Asterichnites octoradiatus. The presence of these trace fossils can aid in understanding
vertebrate behavior and the paleoecology of the Mowry Sea, as well as provide insight into the paleoenvironmental conditions
of the Cretaceous marine ecosystems of the Western Interior.

INTRODUCTION Wyoming and to describe new discoveries. The well-

preserved trace fossils provide evidence of behavior of
The Cretaceous Mowry Shale Formation was various vertebrates and cephalopods. Study of the
deposited in an epicontinental sea, covering much of sediments, along with description of the trace fossils,
the Western Interior of North America. The sea can provide a picture of the fauna interacting with their
extended along the east side of the Cordillera, as far environment, and can increase the understanding of
south as Utah and Colorado, but did not connect to the ecosystems of the Mowry Sea.
Tethyan Ocean to the south (Bremer, 2016; Plint et al.,
2009). The Mowry Sea supported a variety of marine STRATIGRAPHY
reptiles including ichthyosaurs (Nace, 1939;
McGowan, 1972; Maxwell and Kear, 2010), At the study sites, the Mowry Formation contains
plesiosaurs, and crocodiles (Massare and Dain, 1986; thinly laminated, dark gray siliceous shale that
Massare, 1998; Stewart et al., 1994). Invertebrates are weathers to a silver-grey. Unlike the shales above and
known mainly from trace fossils. Clark (2010) below the formation, it often hosts small pine forests,
described the ichnofossil Planolites from near the which take advantage of the hard siliceous shale as a
uppermost layer of the Mowry Shale at Alkali water reservoir. This feature makes it easier to find the
Anticline in the Bighorn Basin, near one of the study formation in the field. The Mowry Shale Formation
sites. Invertebrate trace fossils from other localities includes a shale interval from which the formation gets
include Protovirgularia, Cylindrichnus, Lockeia, its name, and the overlying, and much thinner layer,
Planolites, navichnia and Zoophycos (Clark, 2010). the Clay Spur bentonite. Stratigraphically the study
Although trace fossils are known from the Mowry sites are located at the topmost layer of the Mowry
Shale, they are uncommon (Clark, 2010; pers. obs.). Shale and just below a sharp contact with the Clay
Trace fossils are important indicators of past Spur bentonite, the uppermost bed of the Mowry Shale
animal behavior and can be applied to understand the Formation (Figure 1). A hiatus between the Mowry
activity of extinct species (Bennett et al., 2014). How Shale and the Clay Spur bentonite is not present,
an animal interacts with their environment such as the indicated by the clarity and detailed preservation of the
sea floor can provide clues to paleoecology (Rhoads, surface traces, which imply the absence of erosion and
1975). Generally, vertebrate traces in a marine a rapid burial. Weathering features indicating subaerial
ecosystem are rare (e.g., Manni et al., 1999; Zhang et exposure such as desiccation cracks are not present.
al., 2014). At the study sites, however, vertebrate traces The Mowry Shale Formation marks the transition
were the dominant ichnofossil. Those described in this between the Lower and Upper Cretaceous
study are on the top-most surface of the Mowry Shale. (Albian/Cenomanian transition) units in Central
This study is an attempt to investigate known Wyoming. Cobban and Reeside (1951) and Reeside,
ichnofossils localities in the Mowry Shale Formation in and Cobban (1960) placed the Mowry Shale on the

68
 CONNELY—VERTEBRATE TRACE FOSSILS 69

Cenomanian/Albian boundary based on ammonite and Vokes (1944) as a cephalopod trace fossil,
assemblages. Based on biostratigraphic and sequence Asterichnities octoradiatus. However, Vokes (1941, p.
stratigraphic correlations of North Texas and European 452) also showed an image he described as a
sections, along with radiometric dating of bentonite “…groove-like gastropod (?) trail,…” These trails
beds in the Western Interior sediments of North have been noted in other areas and are the focus of this
America, the Albian/Cenomanian boundary has been report. Davis (1963, p. 143), mentions “…long,
set at 97.2 Ma (Scott et al., 2009). However, the sinuous, groove-like depressions about one-half to one
International Commission on Stratigraphy (2018) inch across, one-half inch deep, and up to sixty feet
currently places the boundary at 100.5 Ma. Researchers long wind across this bedding plane surface…” in the
radiometrically dated the “Clay Spur” bentonite at ~98 upper contact of the Mowry Shale from Weston
Ma (Obradovich et al., 2002; Obradovich, 2005, pers. County, Wyoming. Burford (1985) described similar
comm,), making it Cenomanian in age. George et al. markings at a site located in the Bureau of Land
(2014) placed the boundary between the Upper and Management’s (BLM) Off Highway Vehicles (OHV)
Lower Cretaceous at the base of the Mowry Shale Park in Natrona County, Wyoming (Hanson and
(Figure 1), making all of the Mowry Shale Formation Connely, 2006). Burford (1985) mentioned that these
Cenomanian in age. trace fossils could possibly belong to a marine
Overlying the Clay Spur bentonite is the Frontier crocodile.
Formation (Watson, 1980) west of the Powder River Some of the trace fossils mentioned by Burford
Basin, and the Belle Fourche Shale (Hosterman and (1985) have now been re-described as belonging to the
Patterson, 1992; Massare, 1998) east of the basin, both ichnotaxa Asterichnites octoradiatus (Figure 2).
of which are Cenomanian in age (George et al., 2014; However, the other trace fossils from the OHV Park
International Chronostratigraphic Chart, 2018; Figure were not identified. In 2004, trace fossils from the
1). The contact between the Mowry Shale Formation OHV Park were reported to the Tate Geological
and the Frontier Formation is typically placed at the top Museum by local citizens. Upon further investigation,
of the Clay Spur bentonite bed (Nixon, 1973; Burtner it was determined that this was the same stratigraphic
and Warner, 1984). Underlying the Mowry Shale is the interval as those reported from Johnson County (Figure
Muddy Sandstone, which is Albian in age (George et 3) and in Big Horn County, in an outcrop of the Alkali
al., 2014; International Chronostratigraphic Chart, Anticline in the Bighorn Basin (i.e. uppermost layers of
2018). the Mowry Shale). These reports were presented as
____________________________________________ undergraduate research projects (Connely and Talbot,
2005; Trumbull and Connely, 2010).

FIGURE 1. Stratigraphic chart of Cretaceous units associated with

the Natrona County, Wyoming. Taken in part from the Wyoming
Stratigraphic Nomenclature Chart (George et al., 2014).
_______________________________________________________
FIGURE 2. Asterichnites octoradiatus trace fossils at the OHV Park,
Natrona County, Wyoming. Scale = 15 cm.
LOCALITIES WITH TRACE FOSSILS

History of the Sites.—Vokes (1941) first

reported trace fossils of unknown origin in the Mowry
Shale. These were later described and named by Brown
70 PALUDICOLA, VOL. 12, NO. 2, 2019

FIGURE 3. Trace fossils reported from Johnson County, Wyoming by Lawson (2009). This site did not have the protective “sandpaper” surface and
was destroyed by weather shortly after this image was taken. Notice several parallel grooves and one single groove cutting across from lower left to
upper right.
________________________________________________________________________________________________________________________

Location of the Sites.—A total of six sites the eastern flank of a northwest, southeast trending
throughout Wyoming and surrounding states have an structure known as the Immigrant Gap Anticline,
assemblage of trace fossils on the top surface of the northwest of Casper. The surface containing the trace
Mowry Shale, presumably the same stratigraphic level fossils was uncovered by the mining of the Clay Spur
(Figure 4). The site with the most diverse collection is bentonite. This exposed shale surface was protected by
the BLM OHV Park in Natrona County, west of a sandpaper–like layer described as a “crystalline ash”
Casper. The original discovery site containing trace at another nearby locality (Corbett et al., 1985). The
fossils was described by Vokes (1941) from an area surface contains a single layer of light colored, angular,
southwest of Billings, Montana. Brown and Vokes course sand-sized grains mostly of quartz, biotite, and
(1944) described similar tracks to an area northwest of sanadine over a layer 1 to 4 mm thick of similar
Cody, Park County. Davis (1963) reported his material but much finer grained. This single layer of the
observations from Weston County. The exact locations sandpaper surface appears to cover all of the tracks and
of the historic sites are unknown. The fifth site is traces and was likely deposited shortly after the traces
located in Johnson County in an active bentonite mine were made. Fragmentary vertebrate remains, possibly
(Lawson, 2009, pers. comm.). Here numerous trace an ichthyosaur, were present at the surface of the OHV
fossils were reported (Figure 3). However, shortly after Park. Unfortunately they were too weathered for a
the site was identified, the surface was destroyed by positive identification.
winter weather conditions and thus could not be studied
in detail. The sixth and most recently discovered site is Methods.—The study was started in 2004 and
located in the Alkali Anticline in Big Horn County continues to this day. The surface of the outcrops
(Trumbull and Connely, 2010). containing the trace fossils was mapped using a grid
The outcrop that contains the OHV Park site is on system. The data was transferred onto Mylar grid paper
 CONNELY—VERTEBRATE TRACE FOSSILS 71

FIGURE 4. Locations of Wyoming sites reported in this study.

________________________________________________________________________________________________

to illustrate the morphology and orientation of traces. In Fossils are known from the Mowry Shale, but are
some cases, latex molds were made in the field. These fragmentary (Druckenmiller, 2002), including
molds are curated at the Tate Geological Museum, vertebrates, invertebrates, plants, and ichnofossils.
Casper College, Wyoming. Ongoing studies are Although rare, ammonoids and bivalves are the most
incorporating photogrammetry to record trace common invertebrates observed in the Mowry Shale in
morphology. Natrona County. At the upper most surface containing
the trace fossils, ammonoids were present only at the
MOWRY SHALE PALEOENVIRONMENT AND Johnson County site. Vuke (1982) described some well-
FOSSILS preserved angiosperm leaf fossils from the Galalatin
Ranges of Montana, which could be some of the earliest
The Mowry Shale represents high stand deposition examples known. Various pollens and other
(Long et al., 2000) with a depth of 15m to 150m at the microfossils are also known and were used in
OHV Park site about the time that these traces were biostratigraphic studies (Vuke, 1982; Scott et al., 2009).
created. Further west, the maximum depth was 300m However, the stratigraphic position of these reports is
(Costanzo, 2006). The identified ichnofossils unclear.
represented by Zoophycos and Cruziana ichnofacies Vertebrates reported from the Mowry Shale
and the lack of sedimentary structures suggests water include fish such as Hypsodon sp., Leucicthy sp.,
depths beneath fair-weather wave base to beneath Erythrinolepis sp., and Halecodon sp. (Cockerall,
storm-wave base (Bremer, 2016). At the maximum 1919; Anderson and Kowallis, 2005). Most fish are
depths, the water would have been aerobic to identified through abundant scales in the shale.
dysaerobic (Elzea and Murray, 1990). Stewart and Hakel (2006) collected eight types of
72 PALUDICOLA, VOL. 12, NO. 2, 2019

Actinopterygii from the Mowry Shale in Natrona

County. Turtles, crocodiles, and sharks (Paull, 1957)
were found in the underlying Muddy Sandstone, and
marine crocodiles are also known from the Mowry
Shale (Mook, 1934). Plesiosaurs and ichthyosaurs are
well known from the Mowry Shale (Nace, 1939;
Romer, 1968; Massare and Dain, 1989; Stewart, et al,
1994), the latter being the most common marine
reptile in the fauna (J. Massare, 2019, pers. comm.).
At the OHV Park, a partially articulated ichthyosaur
skeleton was collect from a hard ironstone concretion
only a few hundred meters from the trace fossil sites
by the staff of the Tate Geological Museum, Casper
College, Wyoming. Other unidentified bones were
found within the trace layer. Scattered bones of
marine, flying, and even terrestrial reptiles were found
in concretions in the Mowry Shale, but details of the
species have not been described (Kirschbaum and
Roberts, 2005).

DESCRIPTION AND INTERPRETATION

Several fossil tracks, traces, and impressions

were identified at the OHV Park and three other sites
in Wyoming. Five of these morphotypes (herein
referred to as A – E) are most likely from vertebrates,
whereas a few other morphotypes (including the
ichnotaxon Asterichnites octoradiatus) are probably
from invertebrates. FIGURE 5. Trace fossil Type A from the OHV Park, Natrona
County, Wyoming. The surface is protected by the “sandpaper”-like
Vertebrate Traces: Trace Type A.—These trace
surface. Notice the expulsion material along the edge of the trace.
fossils are long (1 to ~26 meters) single or two parallel Scale = 10 cm
grooves, ~3 cm wide and .5 to 1.5 cm deep (Figure 5). _______________________________________________________
The groove vertical section is u-shaped. The sides of
the grooves are slightly raised with an expulsion or main propulsion, whereas plesiosaurs swam using their
displacement rim. Most grooves are straight although limbs. The tail of the plesiosaur or the pectoral fins of
longer sets have a single bend to an s-shaped pattern. the ichthyosaur was used for stability (Massare, 1988).
Parallel sets are ~ 35 to 50 cm apart (Figure 6). The Many of these trace fossils come in parallel pairs which
grooves are not always continuous but become shallow would support a pectoral fin or appendage
for about a meter then reappear along the same vector. interpretation. The groove is long and shallow (U-
These fossils match the descriptions made by Davis shaped vertical section) created by the dragging of a
(1963) from a mine in Weston County. These trace blunt object. Ichthyosaurs have stiff and fairly thick
fossils appear to have been made by a swimming forelimbs. Fish tend to have thinner blade like fins,
organism dragging an appendage in the mud. The which would create a more V-shaped impression.
expulsion rim along the edges supports a displacement Crocodiles and some fish are more undulatory in their
of mud or marginal ridge and is not a depositional swimming movement, using much of their torso for
artifact. The little (or lack of) sinuosity suggests that the propulsion (Massare, 1988). This type of movement
track maker used an axial oscillatory or suboscillatory would show up in a sinuous pattern (such as the
mode of movement. This appendage or tail is likely not inchotaxon Undichna Anderson, 1976). Tool marks can
responsible for the primary propulsion of the animal but be ruled out because these grooves are numerous,
may have been used to steer or stabilize the body. The crisscross each other with no consistent orientation, and
gradual skips in the trace suggest that the track maker do not vary their morphology, size and distance
periodically became more buoyant as it was moving between each other along their course.
through the water column, and was not a benthic Trace Type B.—These trace fossils are two
creature. These traces might have been created by the separate but apparently related markings (Figures 7, 8
pectoral fin of an ichthyosaur or the tail of a plesiosaur. and 9). The primary trace has a broken, sinuous pattern
Ichthyosaurs have a lunate tail, which is used for their or arcuate impression. They are thinner in width (~1cm
 CONNELY—VERTEBRATE TRACE FOSSILS 73

related as they occur together in both instances,

however they are offset and do not overlap like those in
Undichna. This trace fossil is interpreted as made by a
single individual with a tail, which uses oscillatory or
suboscillatory propulsion. Aquatic amphibians with a
long tail and trailing legs (Turek, 1989) as well as
aquatic reptiles (Trewin, 2000) could make similar
impressions. Swimming crocodiles use their tail with
suboscillatory propulsion (Massare, 1988) or faster
axial swimming (Farlow, et al., 2018). Modern
crocodiles use its hind limbs for balance, steering and to
push off of the bottom or paddle the animal forward
(Farlow, et al., 2018). This posture would allow claws
to gently mark the sea floor. However, it is unlikely that
dragging claws could make even parallel marks like
those described in Trace Type B. With some
exceptions, the trace fossil best resembles that of
Undichna insolentia as described by Trewin (2000). U.
insolentia displays four parallel in-phase and incised
epichnial grooves. Trace Type B differs from U.
insolentia in that the inner pair of the parallel grooves
are not strongly incised and is twice the size as
Undichna described by Trewin (2000). In addition,
Undichna is associated with oxygenated freshwater
environments instead of the more reducing, marine
environment of the Mowry Shale.
Trace Type C.—These trace fossils are shallow
(~1 cm deep) and longer than wide (25 cm wide and 1
to 5 meters long) troughs (Figure 10, 11, and 12). Each
FIGURE 6. Two parallel traces of Type A from the OHV Park. trace fossil has a gradual end with no apparent edge,
_____________________________________________ whereas the other end may have a well-defined
terminus. All appear to be impressed in a north/south
wide) and shallower in depth (<1cm) then those in Type direction at the OHV Park. Some have a subtle
A. The overall length is much shorter and with a secondary central indentation in a shallow ridge down
repeated sinuous morphology. In cross section, these the center. The surface of the trace fossil is smooth but
traces are U to V-shaped. The secondary traces are thin with uneven sides. A marginal ridge is faint and non-
3 to 4 parallel markings that can be found in a existent in many samples. This track was not identified
subparallel path to the primary trace. These have a at the Johnson County or the Alkali Anticline sites in
similar sinuous pattern but are more continuous. This Big Horn County. These may be similar to a second
trace fossil was only observed at the OHV Park, where type of trace fossil described by Davis (1963) in
two occurrences were mapped. Both trace fossils can be Weston County.
compared to the fish swimming trace ichnogenus Type C trace fossil appear to be more of a partial
Undichna (Anderson, 1976). They also resemble the resting trace than that of a swimming trace. The mud is
crocodile swim trace fossil (Hatcherichnus) from the slightly disturbed as an indentation. Expulsion material
Upper Jurassic Morrison Formation (Foster and is not present. The trace fossil resembles the impression
Lockley, 1997) but without the manus or pes made by an animal coming down to rest on the ocean
impressions. bottom and coming to a complete stop as it may be
These trace fossils suggest an animal swimming inferred by the terminus end and a shallow beginning. It
near the seafloor. The primary trace appears to be is unclear as to the trace maker, but it is clearly made
produced by a tail/fin that was propelling a swimming by an animal that has a flat bottomed or slightly convex
animal with axial horizontal oscillatory or torso, with a hint of a medial furrow, and is
suboscillatory motion. The secondary parallel grooves approximately 20 to 25 cm wide. If these represent a
on the side are more likely an appendage mark resting trace fossil, an ambush predator would be a
generated by claws, sharp spines, or fins that drag or cut
through the mud. The two trace fossils seem to be
74 PALUDICOLA, VOL. 12, NO. 2, 2019

FIGURE 7. Trace fossil Type B. Primary trace with line drawing from OHV Park, Natrona County, Wyoming. Scale = 15 cm.
___________________________________________________________________________________________________ _______

FIGURE 8. Trace fossil Type B. Secondary trace with line drawing. Scale = 15 cm.
 CONNELY—VERTEBRATE TRACE FOSSILS 75

FIGURE 9. Trace fossil Type B with associated primary and secondary trace set.
_________________________________________________________________________________________________________________

FIGURE 10. Trace fossil Type C. Resting trace with no expulsion rim.
76 PALUDICOLA, VOL. 12, NO. 2, 2019

FIGURE 11. Trace fossil Type C showing terminus end (upper right). Scale = 15 cm.
____________________________________________________________________________________________________________________

likely candidate. Crocodiles, which fit the size of the Trace Type D.—These trace fossils are
trace fossil, are described as ambush predators indentations showing possibly 2 or 3 impressions
(Massare, 1988) and could easily rest on the ocean floor (Figure 13). There is only one example of this trace so
to wait for food to pass by. Forrest (2003) described any interpretation is highly speculative. It could be an
evidence from a plesiosaur that had been partially eaten invertebrate resting trace fossil such as Rusophycus.
by the crocodile Metriorhynchus showing related Trace Type E.—These trace fossils are
behavior. Modern species are able to submerge for up irregularly sub-circular impressions (Figure 14). In
to two hours (Seebacher, et al., 2005) and at depths of cross section one edge is deeper and becomes shallow
several meters. Furthermore, much of their social, at the other end. Just outside of the deep end side are
feeding and reproductive behavior often occurs in the claw-like marks and a pile of ejecta. There are only two
water (Farlow et al., 2018). As mentioned by Forrest of these trace fossils, which appear to be associated and
(2003), crocodiles are capable of exploiting a wide oriented in the same direction. These trace fossils
range of feeding sources and habitats. Other animals resemble descriptions by Farlow et al. (2018) and
that could leave this type of impression would include Natali et al. (2019).
large fish, sharks, and turtles. Invertebrates are ruled This trace fossil is only known from the OHV
out due to the size of the trace fossil. Although Park. However, fossil trackways of a possible dinosaur
ammonites are known from the Mowry Shale were reported from the Mowry Shale on Dinosaur
Formation, most are too small to create a 25 cm wide Ridge, Colorado (Moklestad et al., 2018). The deep
shallow trace fossil. The lack of fin marks and general impression, claw-like marks, and ejecta from Type E
size rule out most fish types. The sediments appear to suggest it was made by a clawed animal pushing off of
be displaced but not overly disturbed. Therefore, it is the ocean bottom or a bottom walking and/or punting
doubtful that this is a feeding behavior of an animal reptiles. Crocodiles and turtles are the only marine
looking for food within the mud of the ocean floor. animals with claws. Crocodiles and turtles are known
 CONNELY—VERTEBRATE TRACE FOSSILS 77

Other trace fossils observed at the site include a surface

feeding trace resembling that of an unbranched or semi-
spiral burrow. Vertical trace fossils were also seen but
due to the laminar fracturing pattern of the Mowry
Shale, finding an intact burrow was difficult and will be
investigated in future studies.
Asterichnites octoradiatus (Brown and Vokes,
1944, pg 658) was found along with the vertebrate
traces described above (Figure 2). These are the most
common invertebrate traces in the study areas.
Asterichnites octoradiatus are eight-limbed star-like
impressions. Each has a center circular quarter-sized
impression with eight thin grooves radiating away from
the center. Brown and Vokes (1944) described these as
being possible cephalopod impressions. They are the
most common trace for all of the studied sites, but this
may be due to the unique morphology that makes them
easily recognizable. Upon careful inspection of one
sample (TATE i419), a second set of parallel grooves
inside each tentacle impression could be seen (Figure
15). The presence of the secondary grooves within the
tentacle impressions could easily have been created by
the hooklets found on the tentacles of a squid-like
animal.

FIGURE 12. Trace fossil Type C showing medial ridge. Scale = 15

cm.
________________________________________________________

bottom walkers (Brand, 1979; Bennett et al., 2014;

Xing et al., 2014; Farlow et al., 2018) and could have
left this type of track morphology. Turtles are known
from the underlying Muddy Sandstone, but not from the
Mowry Shale, so they are a less likely candidate for the
trace fossil maker. Crocodile swim traces are also
known from another Cretaceous deposit, such as the
Dakota Group of Colorado (Foster and Lockley, 1997;
Avanzini et al., 2007; Lockley et al., 2010), and is a
more likely candidate. Natali (2019) described similar
traces in marine sediments of the Maiolica Formation in
Italy. He specifically described tetrapod tracks that are
irregularly sub-circular or elliptical shape with a
displacement rim, somewhat similar to Type E. Natali
(2019) suggested that these traces were made by the
forelimb based on the lack of an overprint. Trace Type
E appears to only represent only one side of the body if
they are consecutive impressions. This feature is not
uncommon, as was reported by Farlow et al., (2018).

Invertebrate Traces.—A few invertebrate trace fossils

were found on the same surface as the vertebrate trace
fossils. Invertebrates trace fossils are not as common at
the study sites, with the exception of the supposed FIGURE 13. Trace fossil Type D. Scale = ~ 10 cm.
cephalopod trace fossil (see below). Planolites was
described near the Alkali Anticline site (Clark, 2010).
78 PALUDICOLA, VOL. 12, NO. 2, 2019

FIGURE 14. Trace fossil Type E showing mud drops or ejecta (left) and track impression on the right. Scale = 15 cm.
______________________________________________________________________________________________________________

A
B

FIGURE 15 Asterichnites octoradiatus (TATE i419). A.Close up view showing secondary grooves. B. Details of groove and hooklet impressions.
Scale in mm.
 CONNELY—VERTEBRATE TRACE FOSSILS 79

FIGURE 16. Illustration showing ecosystem interpretation. © Copyright Russell Hawley 2019.

The behavior associated with these trace fossils is cephalopods, then it wouldn’t be hard to assume that
unclear. Brown and Vokes (1944) suggested that the marine reptiles would find the ocean bottom a good
cephalopod was possibly looking for a place to attach place to find a meal (Figure 16). The preserved
eggs. During reproductive activity, squid deposit egg stomach contents of marine reptiles (Pollard, 1968;
sacks on the ocean floor. In modern squid, these Massare and Young, 2005) suggest that squid and other
deposits can be communal in a “big bang” deposit or cephalopods were a common food source.
they could be singular solitary deposits (Hanlon and
Messenger, 1996). Furthermore, some squid are known CONCLUSION
to migrate to shallow waters to spawn, although others
will stay in deeper waters. The tracks do not seem to be The uppermost surface of the Mowry Shale in
random but often occur as ~10 impressions in a semi- Wyoming provides researchers with well-preserved set
linear pattern. This would suggest an intentional of trace fossils from Cretaceous (Cenomanian) marine
behavior such as in egg laying. Drew (1911) recorded a ecosystem. Ash from volcanic eruptions buried these
female Doryteuthis münster bouncing along the ocean traces, capturing a single moment in time. It appears
bottom with only the tips of her arms just before that the Mowry had a diverse fauna of predators and
depositing an egg string. The trace fossils at the OHV prey. The trace fossils found on the ocean floor show
Park and those described by Vokes (1941) have rows of evidence of behavior from various vertebrates and
stellate impressions and are arranged in a semi-linear cephalopods, although interpretations are admittedly
pattern, which would support the bouncing hypothesis. subjective. Continued studies and new discoveries will
A second possibility would be a feeding behavior, help clarify these explanations.
where the cephalopod would probe the ocean floor for The current interpretation describes a scene
invertebrates. The presence of various worm burrows where predators, both ambush predators and active
in the shale, although uncommon, would support this hunters, rest and scour the ocean bottom looking for
idea. However, it seems that this behavior would prey (Figure 16). Jurassic ichthyosaurs fed on
produce a more random pattern in the track positions cephalopods, as indicated by hooklets found as stomach
on the ocean floor. In either case, if this site was a contents (e.g., Pollard, 1968; Massare and Young,
common place for egg laying or a feeding ground for 2005) although some Cretaceous species had a different
80 PALUDICOLA, VOL. 12, NO. 2, 2019

diet (Kear, et al., 2003). The presence of the Coconino Sandstone (Permian) vertebrate foot-
cephalopod stellate impressions along with evidence of prints and their paleoecological implications.
cruising behavior of ichthyosaurs, as demonstrated by Palaeogeography, Palaeoclimatology, Palaeo-
Trace Type A and B, paints a picture of swimming ecology 28:25-38.
ichthyosaurs and/or other vertebrates looking for Bremer, J.M. 2016. Stratigraphy and sedimentology of
cephalopod and/or their eggs on the seafloor. Trace the Cretaceous Mowry Shale in the Northern
fossil Types C, and E suggest a possible ambush Bighorn Basin of Wyoming: Implications for
predator resting and/or pushing off the sea floor perhaps unconventional resource exploration and
waiting for a fish or other vertebrate to swim by. development. Unpublished M.S Thesis,
University of Nebraska, Lincoln 82 pp.
Brown, B. and H.E. Vokes. 1944. Fossil imprints of
ACKNOWLEDGMENTS unknown origin; 2 Further information and a
possible explanation. American Journal of
This project is dedicated to Arthur E. Burford, a Science 242:656-672.
professor for the University of Akron and a long time Burford, A.E. 1985. Reptilian markings on the Upper
Wyoming geologist. Mr. Burford was contacted in the Mowry Shale Emigrant Gap area, Natrona
summer of 2004 for information about the location of County, Wyoming. The Cretaceous Geology of
the OHV Park site. Unfortunately, Mr. Burford passed Wyoming, Wyoming Geological Association
away a month later before we could tell him about the 36th Annual Field Conference Guidebook 157-
finds and project that has generated from his original 158.
report in 1985. Burtner, R.L. and M.A. Warner. 1986. Relationship
Thanks is given for the following: The NSF between illite/smectite diagenesis and
EPSCoR for funding this project; the Bureau of Land hydrocarbon generation in Lower Cretaceous
Management for their financial and academic support Mowry and Skull Creek Shales of the Northern
and for providing field assistance in mapping a Rocky Mountain Area. Clays and Clay Minerals
surveying; the Tate Geological Museum for access to 34:390-402.
the collections and as the repository for the data and Clark, C.K. 2010. Stratigraphy, sedimentology, and
samples collected; to Russell Hawley for the wonderful ichnology of the Upper Cretaceous Frontier
illustrations and reconstruction of the Mowry Sea; and Formation in the Alkali Anticline region,
to Dr. Obradovich for supplying updated radiometric Bighorn County, Wyoming. Unpublished M.S.
dates. Thesis, University of Nebraska, Lincoln 76 pp.
Cobban, W.A., and J.B. Reeside Jr. 1951. Lower
LITERATURE CITED Cretaceous ammonite in Colorado, Wyoming
and Montana. America Association of Petroleum
Anderson, A. 1976. Fish trails from the Eargly Permian Geologists Bulletin 35:1892-1893.
of South Africa. Palaeontology 19: 397-409. Cockerell, T. D. A. 1919. Some American Cretaceous
Anderson, A.D., and B.J. Kowallis. 2005. Storm fish scales. United States Geological Survey
deposited fish debris in the Cretaceous Mowry Professional Paper 120I:165–188.
Shale near Vernal, Utah. Pp. 125-130, in C.M. Connely, M.V. and B. Talbot. 2005. Trace fossils in
Dehler, J.L.Pederson, D.A.Sprinkel, and B.J. marine sediments of the Mowry Shale (Early
Kowallis (eds.), Uinta Mountain Geology, Utah Cretaceous) of Wyoming are possibly vertebrate
Geological Association Publication 33. in origin. Journal of Vertebrate Paleontology,
Avanzini, M., L. Piñuela, J. Ignacio Ruiz-Omeñaca, and Supplement 25:47.
J. Carlos Garcia-Ramos. 2007. The Crocodile Corbett, R.G., Friberg. L.M., Muller, A.J., Wingard,
Track Hatcherichnus, From the Upper Jurassic P.S. 1985. Sandpaper surface at the superface of
of Asturias (Spain). Pp. 88-92, in J.Milàn, S.G. the Mowry Shale, Natrona County, Wyoming.
Lucas, M.G. Lockley, and J.A. Spielmann, Geological Society of America, Rocky Mountain
(eds.), Crocodyle tracks and traces. New Mexico Section Meeting, Abstracts 17(4):214.
Museum of Natural History and Science, Costanzo, L.A. 2006. Stratigraphy of the Lower
Bulletin 51. Cretaceous Muddy Sandstone, Wind River
Bennett, M.R., S.A. Morse, and P.L. Falkingham. 2014. Indian Reservation, Fremont County, Wyoming.
Tracks made by swimming Hippopotami: An Unpublished M.S Thesis, Colorado School of
example from Koobi For a (Turkana Basin, Mines 151 pp.
Kenya), Palaeogeography, Palaeoclimatology, Davis, J.C. 1963. Origin of the Mowry Shale.
Palaeoecology 409: 9–23. University of Wyoming, Contributions to
Brand, L. 1979. Field and laboratory studies on the Geology 2:135-146.
 CONNELY—VERTEBRATE TRACE FOSSILS 81

Drew, G.A. 1911. Sexual activities of the squid: Loigo Lockley M.G., S.G. Lucas, J. Milàn, J.D. Harris, M.
pealii (Les.): I copulation, egg laying, and Avanzini, J.R. Foster, and J.A. Spielmann. 2010.
fertilization. Journal of Morphology 22:327-352. The fossil record of crocodylian tracks and
Druckenmiller, P.S. 2002. Osteology of a new traces: an overview. Pp. 1-13, in J. Milàn, S.G.
plesiosaur from the lower Cretaceous (Albian) Lucas, M.G. Lockley, and J.A. Spielmann (eds.),
Thermopolis Shale of Montana. Journal of Crocodyle tracks and traces. New Mexico
Vertebrate Paleontology 22:29–42. Museum of Natural History and Science Bulletin
Elzea, J.M., and H. H. Murray. 1990. Variation in the 51.
mineralogical, chemical and physical properties Long, M.S., S.M. Ritter, and T.H. Morris. 2000. Facies
of the Cretaceous Clay Spur bentonite in architecture and sequence stratigraphy of the
Wyoming and Montana: Applied Clay Science Lower Cretaceous Muddy Formation in the
5:229-248. southeastern Bighorn Basin, Wyoming:
Farlow, J.O., N.J. Robinson, M.L.Turner, J. Black, and American Association of Petroleum Geology,
S.M. Gatesy. 2018. Footfall Pattern of a Bottom- Annual Meeting, Abstracts 2000:87.
Walking Crocodile (Crocodylus acutus). Palaios Manni, R., U. Nicosia, and G. Nobili. 1999. An unusual
33(9), 406-413. tetrapod trackway from lower Jurassic marine
Forrest, R. 2003. Evidence for scavenging by the sediments of central italy: Accordiichnus natans
marine crocodile Metriorhynchus on the carcass n. ichnogen., n. ichsp. Geologica Romana
of a plesiosaur. Proceedings of the Geologists 35:167-187.
Association 114:363-366. Massare J.A., 1988. Swimming capabilities of
Foster, J.R., and M.G. Lockley. 1997. Probable Mesozoic marine reptiles: implications for
crocodilian tracks and traces from the Morrison method of predation. Paleobiology14:187-205.
Formation (Upper Jurassic) of eastern Utah: Massare, J.A. 1998. Marine reptiles of the Mowry and
Ichnos 5:121–129. Belle Fourche Shales (Cretaceous) in
George, L.,D. Cardinal, and G. Winter. 2014. Wyoming Northeastern Wyoming. Pp. 33, in R.A. Hunter
Stratigraphic Nomenclature Chart. Wyoming (ed.), Life in The Late Cretaceous, Tate
Geological Association. Geological Museum.
Hanlon, R.T. and J.B. Messenger. 1996. Cephalopod Massare, J.A., and L. E. Dain. 1989. The marine
Behavior. Cambridge University Press, 232 pp. reptiles of the Mowry Shale (Albian) of
Hanson, D.A., and M. Connely. 2006. Mowry Shale northeastern Wyoming. Journal of Vertebrate
ichnofossils; Management of a unique fossil Paleontology. 9(3 supplement): 32A.
tracksite in an off-highway vehicle recreation Massare, J.A. and H.A. Young. 2005. Gastric contents
park. New Mexico Museum of Natural History of an ichthyosaur from the Sundance Formation
and Science, Bulletin 34:19. (Jurassic) of Central Wyoming. Paludicola 5:20-
Hosterman, J.W., and S.H. Patterson. 1992. Bentonite 27
and Fuller’s Earth Resources of the United Maxwell, E.E. and B.P. Kear 2010. Postcranial
States. U. S. Geological Survey Professional anatomy of Platypterygius americanus (Reptilia:
Paper 1522:45pp. Ichthyosauria) from the Cretaceous of Wyoming.
International Commission on Stratigraphy. 2018. Journal of Vertebrate Paleontology 30:1059–
International Chronostratigraphic Chart, 1068.
Available online at www.stratigraphy.org/ McGowan, C. 1972. The systematics of Cretaceous
ICSchart/ChronostratChart2018-07.pdf. ichthyosaurs with particular reference to the
Kear, B.P., W.E. Boles, and E.T. Smith. 2003. Unusual material from North America. University of
gut contents in a Cretaceous ichthyosaur. Wyoming, Contributions to Geology 11:9–29.
Proceedings of the Royal Society of London. Moklestad, T., T. Caneer, and S. Lucas. 2018. The
Suppl 2. Series B: Biological Science 270:206- “Lost Tracks” at Dinosaur Ridge, Colorado,
208. from the base of the Cretaceous (Late Albian-
Kirschbaum, M., and L.N.R. Roberts. 2005. Early Cenomanian) Mowry Shale Member of the
Stratigraphic framework of the Cretaceous Mow- Benton Formation, show a swimming(?)
ry Shale, Frontier Formation and adjacent units, ornithopod affected by a current. Pp. 503-511 in
southwestern Wyoming Province, Wyoming, S.G. Lucas, and R.M. Sullivan, (eds.) Fossil
Colorado, and Utah. Chapter 15, in Petroleum Record 6. New Mexico Museum of Natural
Systems and Geologic Assessment of Oil and History and Science Bulletin 79.
Gas in the Southwestern Wyoming Province, Mook, C.C. 1934. A new species of Teleorhinus from
Wyoming, Colorado, and Utah. U.S. Geological the Benton shales. American Museum Novitates
Survey Digital Data Series DDS–69–D. 702:1-11.
82 PALUDICOLA, VOL. 12, NO. 2, 2019

Nace, R.L. 1939. A new ichthyosaur from the Upper Stewart, J.D., S.A. Bilbey, D.J. Chure, S.K. Madsen,
Cretaceous Mowry Formation of Wyoming. and K. Padian. 1994. Vertebrate fauna of the
American Journal of Science 237:673–686 Mowry Shale (Cenomanian) in northeastern
Natali, L., A. Blasetti, and G., Crocetti. 2019. Detection Utah. Journal of Vertebrate Paleontology 14 (3
of Lower Cretaceous fossils impressions of a supplement):47A.
marine tetrapod on Monte Conero (Central Italy). Stewart, J.D., and M. Hakel. Ichthyofauna of the
Cretaceous Research 93:143-150. Mowry Shale (early Cenomanian) of Wyoming.
Nixon, R.P. 1973. Oil source beds in Cretaceous New Mexico Museum of Natural History and
Mowry Shale of northwestern interior United Science, Bulletin 35:161-163.
States. American Association of Petroleum Trewin, N.H. 2000. The Ichnogenus Undichna, with
Geologists Bulletin 57:136-161. examples from the Permian of the Falkland
Obradovich, J., T. Matsumoto, and T. Nishida. 2002. Islands. Palaeontology 43:979-997.
Integrated biostratigraphic and radiometric study Trumbull, C.B. and M.V. Connely. 2010. New
on the Lower Cenomanian (Cretaceous) of ichnotaxa from the Mowry Shale of Wyoming
Hokkaido, Japan. Proceedings of the Japan may be vertebrate in origin. Geological Society
Academy: Series B. Physical and Biological of America, Rocky Mountain Section Abstracts
Sciences. 78:149–153. 42(3):41-42.
Paull, R.A. 1957. Depositional history of the Muddy Turek, V. 1989. Fish and amphibian trace fossils from
Sandstone Bighorn Basin, Wyoming. Westphalian sediments of Bohemia. Palaeont-
Unpublished Ph.D. Dissertation, University of ology 32:623-643.
Wisconsin. 269 p. Vokes, H.E. 1941. Fossil imprints of unknown origin.
Plint, A.G., A. Tyagi, M.J. Hay, B.L. Varban, H. American Journal of Science 239:451-453.
Zhang, and X. Roca. 2009. Clinoforms, Vuke, S.M. 1982. Depositional Environments of the
paleobathymetry, and ud dispersal across the Cretaceous Thermopolis, Muddy, and Mowry
western Canada, Cretaceous Foreland Basin: Formations Southern Madison and Gallatin
Evidence from the Cenomanian Dunvegan Ranges, Montana. Unpublished M.S. Thesis,
Formation and contiguous strata. Journal of Indiana University. 153 pp.
Sedimentary Research. 79:144–161. Watson, J.E. 1980. Catalog of Wyoming stratigraphy,
Pollard, J. E. 1968. The gastric contents of an Tooke Engineering, Casper, WY
ichthyosaur from the lower Lias of Lyme Regis, Xing, L., M. Avanzini, M.G. Lockley, T. Miyashita, H.
Dorset. Palaeontology 11: 376-388. Klein, J. Zhang, Q. He, L. Qi, J. D. Divay, and
Reeside, J.B., Jr. and W.A. Cobban. 1960. Studies of C. Jia. 2014. Early Cretaceous turtle tracks and
the Mowry Shale (Cretaceous) and contemporary skeletons from the Junggar basin, Xinjiang,
formations in the United States and Canada: A China. Palaios 29:137-144.
stratigraphic and paleontologic study of five Zhang, Q., W. Wen, S. Hu, M.J. Benton, C. Zhou, T.
large assemblages of gastroplitine cephalopods Xie, T. Lü, J. Huang, B. Choo, Z. Q. Chen, J.
interpreted as five species of Neogastroplites Liu, and Q. Zhang. 2014. Nothosaur foraging
having an unusual degree of variability. tracks from the middle Triassic of Southwestern
Geological Survey Professional Paper. 355:1- China. Nature Communications 5, Article
126. 3973:1-12.
Rhoads D.C. 1975. The paleoecological and environ-
mental significance of trace fossils. Pp. 147-160,
in R. W. Frey (ed.) The Study of Trace Fossils.
Springer, Berlin, Heidelberg.
Romer, A.S. 1968. An ichthyosaur skull from the
Cretaceous of Wyoming. University of
Wyoming, Contributions to Geology 7:27–41.
Scott, R.W., F.E. Oboh-Ikuenobe, D.G. Benson and J.
M. Holbrook. 2009. Numerical age calibration of
the Albian/Cenomanian boundary. Stratigraphy
6:17-32.
Seebacher, F., C.E. Franklin, and M. Read. 2005.
Diving behavior of a reptile (Crocodylus
johnstoni) in the wild: Interactions with heart
rate and body temperature. Physiological and
Biochemical Zoology 78:1-8.