Ichnos

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Crocodile and Arthropod Tracks from the Late

Paleocene Wannagan Creek Fauna of North
Dakota, USA

Bruce R. Erickson

To cite this article: Bruce R. Erickson (2005) Crocodile and Arthropod Tracks from the Late
Paleocene Wannagan Creek Fauna of North Dakota, USA, Ichnos, 12:4, 303-308, DOI:
10.1080/1042094050031111

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Ichnos, 12:303308, 2005
Copyright
c Taylor & Francis Inc.
ISSN: 1042-0940 print / 1563-5236 online
DOI: 10.1080/1042094050031111
ICHNOLOGIC NOTE
Crocodile and Arthropod Tracks from the Late Paleocene
Wannagan Creek Fauna of North Dakota, USA
Bruce R. Erickson
The Science Museum of Minnesota, St. Paul, Minnesota, USA
Recently discovered tracks in alluvial sediments of the Late
Paleocene Bullion Creek Formation of western North Dakota have
been identified as belonging to the large eusuchian crocodile Bo-
realosuchus formidabilis and an associated small arthropod. They
are described as two new ichnotaxa. The first of these, Borealo-
suchipus hanksi igen. et isp. nov., is based on a partly complete
trackway having fore- and hind-footprints as well as body marks
preserved in hyporelief. These tracks are directly associated with
numerous body fossils including articulated bones which provide
strong implications for their probable origin. The second specimen,
Koupichnium pentapodus isp. nov., is founded on a trackway of a
small invertebrate in which a series of five pairs of footprints are
arranged along a midline drag mark which is continuous through-
out the length of its trackway. This trackway shares features with
other fossils as well as living arthropod trackways in which four
small feet and a larger posterior hind pusher foot are present.
The two described new ichnotaxa represent the first recognized
tracks from the Wannagan Creek local fauna.
Keywords Wannagan Creek, Paleocene ichnofossils, crocodile and
arthropod tracks
INTRODUCTION
Extensive excavations between 1970 and 1996 in the late
Paleocene Bullion Creek Formation in Billings County, North
Dakota, resulted in a major collection of some 8,000 vertebrate
fossils representing some 20 taxa (Erickson, 1976, 1984, 1991,
1999; Holtzman, 1978). The majority of these specimens were
recovered during development of Wannagan Creek Quarry, site
of Fossil Lake Wannagan (Erickson, 1999).
Other than crocodilian coprolites (Sawyer, 1981; Hallgren,
1981, 1987) and a few invertebrate burrows (Melchior and
Erickson, 1979), no trace fossils were found. This dearth of
evidence is attributed to intensive bioturbation primarily caused
by large abundant crocodiles as indicated by the presence of nu-
merous disarticulated skeletons with disassociated and fractured
Address correspondence to Bruce R. Erickson, The Science
Museum of Minnesota, St. Paul, Minnesota 55102, USA. E-mail:
berickson@smm.org.
303
bones (Sawyer and Erickson, 1998). Beach wash of the shallow
water shoreline sediments of the lake also prevented track preser-
vation. Completion of the lengthy quarry excavations in 1996
initiated an intensive search for additional paleoenvironmental
evidence, especially in the backswamp deposits of the area near
Lake Wannagan. Backswamp deposits of the area are a charac-
teristic fluvial facies of the broad flood plain paleoenvironment
of the Bullion Creek Formation (Royse, 1972; Jacob, 1972) and
yielded a number of additional body fossils as well as the first
trackways.
The only other tracks of vertebrates in the Bullion Creek are
those of birds from a locality about five miles northeast of the
present site. These tracks are solitary prints and stratigraphically
lower than the present trackways but may represent Wannagan
Creek taxa (Kihm and Hartman, 1995). The Wannagan Creek
trackways are preserved in a fine-grained, thin, calcareous sand-
stone layer varying in thickness from 1 to 3 cm. The purpose
of this paper is to describe these new trackways and discuss
their implications in association with body fossils. As Sargeant
(1990) suggested, the vertebrate footprints as well as the inverte-
brate tracks are treated in the same fashion and given ichnofossil
names rather than body fossil names.
DEPOSITIONAL ENVIRONMENT
Paleoenvironmental conditions at Fossil Lake Wannagan and
its surroundings were paludal and fluvial followed by a lacus-
trine environment. Much of the organic content of the deposits
was derived from dense vegetation of shrubs, trees, and an ex-
tensive aquatic flora (Melchior, 1976; Melchior and Hall, 1983).
These environments were not unlike those found today in areas
inhabited by breeding crocodile populations.
The basin occupied by the lake was part of a large flood
plain complex that is characterized by flat-lying sand, clay, and
mudstone beds with interstratified lignitic layers (Royse, 1972;
Jacob, 1972). Flood-plain lakes such as Lake Wannagan origi-
nated by flooding when a river broke through its banks and filled
a low-lying depression. Present-day lakes are often formed in
this way (Selley, 1978). The duration of such a lake is uncertain.
With the buildup of sediments within the basin at Wannagan,
304
a basal marl with overlying organic material, clays, and silts
were introduced into the deeper parts of the basin. Shoreline
sands and mudflats that developed in subaqueous areas near the
lake margins were frequented by basking crocodiles. Therefore,
these places afforded the best situations for the preservation of
tracks such as those described here.
SYSTEMATIC ICHNOLOGY
Borealosuchipus hanksi igen. et isp. nov.
Type specimen: Borealosuchipus hanksi.
Etymology: Borealosuchipus for track made by Borealo-
suchus-like crocodile, not necessarily the genus Borealosuchus,
and hanksi after Douglas Hanks who discovered and collected
the type specimen.
Holotype: SMM P2001.9.1 (Science Museum of Minnesota
specimen) section of matrix containing natural casts of trackway.
Horizon: Bullion Creek Formation, Late Paleocene,
Tiffanian 4.
Locality: Sec. 7, T.141N, R.102W, Billings County, North
Dakota, USA.
FIG. 1. (A) Photograph and (B) line drawing of holotype of Borealosuchipus hanksi, SMM P2002.9.1. (M) imprint of manus, (P) imprint of pes, (F) filled-in
foot imprint, (BD) body drag marks, (DT) direction of travel. Note ripple marks on substrate in photograph (Fig. 1A).
B. R. ERICKSON
Description: Natural casts of a trackway showing fore- and
hindfeet and body traces are preserved in hyporelief at the base of
a sandstone layer as described by Basan (1978). The trackway is
present on four separate pieces of calcareous sandstone (Fig. 1).
Only prints of the right fore- and hindfeet are preserved.
Imprints of the right manus and pes (Fig. 1) clearly show the
presence of digits IIV; however, the tracks lack impressions
of the complete digits and are too obscured to provide detailed
measurements of the feet. A maximum length of the manus
imprint is 9.0 cm measured along digit III and the maximum
breadth of the imprint is 8.5 cm. The maximum length of the
pes imprint is 11.5 cm and the maximum breadth is 11.0 cm.
Angles of convergence and divergence of the feet from midline
cannot be measured because there is no midline reference due
to the incompleteness of the specimen.
The trackway shows belly drag marks (Fig. 1). Spacing of
the individual foot tracks is irregular and the trackway lacks a
stride; therefore, a glenoacetabular distance as estimated by
Schult and Farlow (1992) cannot be indicated. Footprint size,
however, suggests that the trackmaker was close in size to a
mounted skeleton of Borealosuchus formidabilis Erickson SMM
 WANNAGAN CREEK FAUNAL TRACKWAYS
P74.29.6, which has a manus maximum length of 18.3 cm and
breadth (measured at the heel) of 7.6 cm, and a pes maximum
length of 34.0 cm and a breadth (measured at the heel) of 11.0 cm.
The glenoacetabular distance of this skeleton is 89 cm and a total
skeletal length of 3.71 m, which is considered average adult size
within the Wannagan Creek crocodile population.
Discussion and Conclusions: Impressions of the left feet
are missing due to the in situ position of the trackway near the
edge of a cliff whereby the impressions of the left feet were lost
downslope to erosion. Impressions interpreted as body traces
(Fig. 1) suggest either a crawling or a basking behavior. The de-
scribed tracks were made at the sediment/water interface where
the sediment was soft (Seilacher, 1978). This is also indicated by
the short-period ripple marks on the substrate (Fig. 1A). Bask-
ing behavior as described by Webb and Monolis (1989) indicates
that during cool weather a crocodile moves onto a stream or lake
bank to bask in the sun and once it becomes too hot it returns to
the water; thus, it may leave a distinctive track with body marks
(see Sarjeant (1975) for discussion of vertebrate tracks and their
interpretation).
A number of putative crocodilian tracks have been reported
(Padian and Olsen, 1984; Lockley and Hunt, 1994; Lockley et al.,
1995; Foster and Lockley, 1997). Most lack direct associations
with crocodilian body fossils and are recognized because of
their occurrence in sediments presumed to have been deposited
in habitats frequented by crocodiles. General track morphology
of the current specimen bears close resemblance to that of living
crocodilians (Zugg, 1974; Webb, 1977; Reineck and Howard,
1978). Various fossil crocodile and other reptilian tracks are dis-
cussed in detail by Demathieu and Haubold (1974). The track-
FIG. 2. (A) Photograph and (B) line drawing of holotype of Koupichnium pentapodus, SMM P2002.3.3. (S) stride, (TD) tail drag mark, (DT) direction of travel,
(E) enlarged section, see Figure 3.
305
way is also in direct association with numerous body fossils of
the eusuchian crocodile Borealosuchus formidabilis. Among the
forms present that were capable of making a trackway of this
type are this crocodile, the small alligatorine Wannaganosuchus
and Champsosaurus. The latter is an unlikely prospect because
of its elongated phalangeal pattern (Erickson, 1972) and the evi-
dence of a probable footprint of a Cretaceous champsosaur from
Colorado (Lockley and Hunt, 1994, 1995), which also indicates
a longer foot. The large Paleocene taxon Champsosaurus gigas
Erickson of the Wannagan Creek assemblage possesses even
more elongate digits (Erickson, 1985). Wannaganosuchus is a
small form (Erickson, 1982a, 1982b). The known specimens re-
garded as adult-size are considerably less than half the size of an
adult Borealosuchus and would not have been capable of produc-
ing the current trackway. It is concluded that the trackway was
produced by a medium-sized individual of a Borealosuchus-like
crocodile and is referred to the new ichnotaxon Borealosuchipus
hanksi.
Ichnogenus Kouphcinium Nopsca 1923
Kouphichnium pentapodus isp. nov.
Type specimen: K. pentapodus.
Etymology: pentapodus for five pairs of feet.
Holotype: SMM P2002.3.3. Sandstone matrix slab 15
18 cm with single trackway in epirelief.
Horizon: Bullion Creek Formation, Late Paleocene,
Tiffanian 4.
Locality: Sec. 7, T.141N, R.102W, Billings County, North
Dakota.
306 B. R. ERICKSON

FIG. 3. Photographic enlargement of section of holotype of Koupichnium pentapodus, SMM P2002.3.3. (15) foot imprints, (TM) tic marks, (AC) angle of
convergence upon long axis of trackway, (AD) angle of divergence from axis of in-line footprints, (TD) tail drag mark, (DT) direction of travel. See Figure 2 for
complete specimen.

Description: An arthropod trackway showing five pairs of
footprints repeated seven times and a midline drag mark. Im-
pressions of four small, discrete, unequally spaced, pod-like feet
are located medial to a tetradactylous print of a fifth, posterior,
bird-like foot. The four anterior feet are numbered 1 to 4 from
front to rear by convention (Manton, 1977). However, it should
be noted that in some arthropods the order of foot placement
does not necessarily reflect the order of the limbs (Brusca and
Brusca, 1990). This variation may be due to overstepping or
missing strides; therefore, correlating limb with footprint in any
arthropod may be uncertain. Impressions of the second, third,
and fourth feet are unequally spaced and line up with one an-
other medial to the fifth foot (Figs. 2, 3). They converge an-
teriorly toward midline of the trackway with the second being
closest to midline. The angle of convergence is 15 (Fig. 3,
AC). The imprint of the first foot is lateral to that of the sec-
ond at 90 (Fig. 3, AD) and with a spacing like the other in-
line feet. This pattern characterizes the new trackway. Each of
the pod-like prints has a tic mark on its anterior edge caused
by foot drag during forward movement as shown in the tracks
of the presumed spider track Octopodichnus (Sadler, 1993).
Impressions made by each set of four feet further resemble
 WANNAGAN CREEK FAUNAL TRACKWAYS
Octopodichnus (Alf, 1968; Sadler, 1993) except the pattern is
distinct.
The print of a fifth pusher foot has three long, narrow pro-
jections, one directed posteriorly, and a shorter curved projec-
tion located on either side and directed medially and laterally.
As with the other footprints, a distinct tic mark is present at
the anterior edge of the fifth foot (Fig. 3, TM).
Maximum breadth of the trackway is 2.8 cm; maximum
length is 18.0 cm. Stride length measured between the first (num-
ber one) footprint of consecutive sets of prints ranges from 2.2 to
2.6 cm. A narrow drag mark which exhibits a v-shaped bottom
is located along midline of the entire trackway. It is straight and
uninterrupted.
Discussion and Conclusions: Undoubtedly many of the in-
vertebrate tracks that have been described are of arthropods such
as insects, spiders, and crustaceans. This is reasonable given that
over half, some 62%, of an estimated total number of 1.4 mil-
lion living species of organisms are arthropods (Wilson, 1992).
Arthropods show a great variation of tracks and produce many
patterns on substrates made by their multiple foot groups that
reveal their many locomotor styles (Brady, 1947; Fisher, 1978).
Ichnofossils from the Wannagan Creek site, other than crocodil-
ian coprolites (Sawyer, 1981; Hallgren, 1984, 1987), include
only a few trails and burrows some of which may belong to
crustaceans (Melchior and Erickson, 1979). The present track-
way represents walking activity of an arthropod, most likely a
small Limulus-like crustacean with five pairs of legs, a special-
ized pusher hind foot and a sharp, rigid telson that habitually
dragged (Hantzschel, 1975). Sand and mud flats that character-
ize the Wannagan Creek paleoenvironment certainly supported
a variety of arthropod inhabitants. Koupichnium pentapodus is
the first of these to be described.
ACKNOWLEDGEMENTS
The author thanks Drs. Martin Lockley and Richard McCrea
for their critical reviews of the manuscript, which greatly im-
proved its content, and Dr. Ronald Pickerill for numerous ed-
itorial notations. I also thank Debbie Schoenholz and Ellen
Holt-Werle for their efforts in completing the illustrations,
typing, and electronic versions of the manuscript and fig-
ures. Tim Ready provided the original copies of all of the
figures.
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