PALAIOS, 2008, v. 23, p.
700–710
Research Note
DOI: 10.2110/palo.2007.p07-082r
A WET INTERDUNE DINOSAUR TRAMPLED SURFACE IN THE JURASSIC NAVAJO SANDSTONE,
COYOTE BUTTES, ARIZONA: RARE PRESERVATION OF MULTIPLE TRACK TYPES AND
TAIL TRACES
WINSTON M. SEILER* and MARJORIE A. CHAN
University of Utah, Department of Geology and Geophysics, Salt Lake City, Utah 84112, USA
e-mail: winstonseiler@yahoo.com
ABSTRACT
A distinctive, disturbed surface with numerous soft-sediment im-
pressions occurs within a wet interdune interval of Jurassic Navajo
Sandstone at the Coyote Buttes along the Arizona-Utah border. These
high-density impressions are interpreted as footprints that comprise
a dinosaur trampled surface. This surface displays an unusual com-
bination of multiple overlapping track types and sizes, distinct to
modified footprint features that include claws and toes and rare tail
traces. The trampled surface covers ⬃3000 m2 with an average den-
sity of ⬃12 impressions/m2 in its main extent. Although modern wa-
ter collection and biofilms typical of weathering potholes or pits are
superimposed on this surface, the primary origin of the impression
features are trace fossil structures formed prior to lithification. Four
criteria distinguish the impressions as vertebrate in origin: (1)
large—up to several tens of centimeters—repeating identifiable foot
morphologies; (2) impression floors surrounded by soft-sediment
marginal ridges; (3) impressions that are rarely flat and are typically
oriented at an angle into the sediment (media) and indicate a clear
direction of travel; and (4) multiple in situ ichnofossils on a moist
interdune surface that resulted in soft-sediment deformation. At least
three ichnogenera—cf. Eubrontes, cf. Anchisauripus, cf. Grallator—
and the tracks attributed to a sauropodomorph appear as regular to
asymmetric penetrations into the media with digitate features, com-
monly accompanied by soft-sediment marginal ridges of displaced
sand preserved in the sandstone. The trampled surface provides
paleoecologic and paleoclimatologic proxies that suggest a pluvial
climate shift likely induced groundwater saturation of an eolian in-
terdune that attracted dinosaurs to the area. The trampled surface
provides valuable data for refining ecologic and climatic sensitivities
recorded in Early Jurassic eolian deposits.
INTRODUCTION
Well-preserved dinosaur tracksites have been described at numerous
localities in the Western United States. Documented tracks typically ex-
hibit a wide range of occurrences, from individual tracks (e.g., Olsen et
al., 1998) to linked trackways (e.g., Lockley et al., 1986) to less distinct
trampled beds (e.g., Costa Da Silva et al., 2007). Such tracksites occur
in many continental formations from the Late Triassic through Late Cre-
taceous, represent a wide range of depositional environments, and provide
valuable paleoenvironmental and paleobiological information (Lockley
and Conrad, 1989). More than 60 tracksites have been documented in the
Lower Jurassic Navajo Sandstone (Faul and Roberts, 1951; Clark and
Fastovsky, 1986; Winkler et al., 1991; Rainforth, 1997; Lockley, 1998;
Hamblin and Foster, 2000; Irmis, 2005; Loope, 2006). Most of these
tracks were originally interpreted to have formed in wet eolian sands,
although Loope (2006) argues that many Navajo Sandstone tracks can be
attributed to a dry-sand origin.
* Corresponding author.
Copyright 䊚 2008, SEPM (Society for Sedimentary Geology)
This paper documents the initial survey of a disturbed bedding surface
in the Navajo Sandstone interpreted as an interdune trampled surface
produced by dinosaurs. The number of impressions—many hundreds—
in this trampled surface compare to other mega-tracksites of the western
United States (Lockley and Hunt, 1995), including those of the large
trackway sites of the Morrison Formation (Lockley et al., 1986). The
unusual surface of this study lends significant climatic and ecologic in-
sight into conditions in western Pangea during the Early Jurassic.
LOCATION, GEOLOGIC SETTING, AND STRATIGRAPHY
The Jurassic Navajo Sandstone is well exposed at the Coyote Buttes
(N36⬚59⬘45⬙ W112⬚00⬘22⬙) along the Utah-Arizona border on the west-
ern margin of the Paria Plateau ⬃50 km west of Page, Arizona (Fig.
1A). The locale is notable for its unique diagenetic coloration of iron
oxide mineralogies (Seiler, 2008); climatically induced, cyclic, eolian
stratification (Chan and Archer, 2000; Loope et al., 2001); dry-season
dinosaur tracks (Loope, 2006); and erosional sculpting by wind (Loope
et al., 2008). The disturbed bedding surface (Fig. 1B) is located in the
central portion of the Bureau of Land Management Coyote Buttes North
Special Permit Area, on the eastern flank of the north trending Top Rock
topographic high. In this paper, following its initial description, the sur-
face is informally referred to as the Top Rock trampled surface.
The Navajo Sandstone and its equivalent deposits occupy a basinal
area of ⬎265,000 km2, though its original extent may have been two and
a half times what is preserved in the geologic record (Kocurek and Dott,
1983; Blakey et al., 1988; Kocurek, 2003). The eolian Navajo Sandstone
preserves the deposits of an ancient erg and is the youngest unit of the
Glen Canyon Group (e.g., Olsen and Galton, 1977; Clark and Fastovsky,
1986; Irmis, 2005). A vast literature provides context for the sedimen-
tology and stratigraphy of the Navajo Sandstone (e.g., Blakey et al., 1988;
Peterson, 1988; Sampson, 1992; Verlander, 1995; Chan et al., 2000; Ei-
senberg, 2003). Deposition is estimated to have occurred at ⬃10⬚ N
latitude on the western edge of the Pangean Supercontinent (Blakey,
2001; Loope et al., 2004); the generalized Jurassic paleoclimate setting
is summarized by Parrish and Peterson (1988) and Parrish (1993).
Locally, the Navajo Sandstone strikes 355⬚ and dips 5⬚E corresponding
to tilting associated with the Laramide uplift of the East Kaibab Mono-
cline. The Navajo Sandstone on the Paria Plateau is ⬃600 m thick (Loope
and Rowe, 2003; Loope, 2006), and exposure exceeds 200 m at the Coy-
ote Buttes. The disturbed bedding plane occurs in a heavily bioturbated
interdune interval of the Navajo Sandstone (Seiler, 2008). Eolian inter-
dune deposits (e.g., Kocurek, 1981) are distinguished by a combination
of factors that include horizontal bedding, sabkha-type laminations, des-
iccation cracks, rhizoliths, a high degree of bioturbation, and association
with a rising water table, lacustrine, or fluvial deposits.
METHODS
The disturbed bedding plane surface is ⬃1 m thick and covers a rel-
atively large area (⬃3000 m2) that can be viewed only at a low, oblique
0883-1351/08/0023-0700/$3.00
PALAIOS WET INTERDUNE DINOSAUR TRAMPLED SURFACE IN JURASSIC NAVAJO SANDSTONE 701

FIGURE 1—Top Rock trampled surface and representative impressions. A) Coyote Buttes study area. Oblique downward, NNW view of the trampled surface outlined by
dashed line. T ⫽ tail trace location. B) Characteristic impression distribution within the central portion of the trampled surface. C) Trackway of three-toed cf. Grallator.
D) Sinuous, curved tail trace straddled by shallow (1–2 cm), circular (7.7 cm diameter) footprints. Note expulsion rim formed by the tail at the upper right (arrow). E)
Synsedimentary soft-sediment ridges upturned around imprints (white arrows); smaller impressions overprinted atop larger impressions (black arrow). F) Impression forming
a possible trackway (into the frame). G) Diagram of photograph D features. Solid line ⫽ known tail trace; dashed line ⫽ inferred tail trace; solid ellipses ⫽ footprints
belonging to tail-imprinting organism; broken ellipses ⫽ footprints not associated with tail trace.
702 SEILER AND CHAN
angle from above (Fig. 1A). Vertical study would have required the use
of ladders or aerial photography, which was not practical given field area
constraints, including flight restrictions over designated wilderness areas
and national monuments. Unlike other trackways that may have several
to dozens of footprint impressions, this particular surface has ⬎1000
individual depressions that exceed our ability to measure each individu-
ally for this pilot study. To document the significance of this surface, ten
field plots (2 m ⫻ 2 m) totaling 40 m2 (Fig. 2) were surveyed in the
main extent (the eastern 60 m ⫽ ⬃1320 m2) of the disturbed surface,
as a preliminary means to assess the occurrence of impressions. Plots
were chosen randomly to avoid bias towards the larger, more unusual, or
visually exciting impressions, with the stipulation that plots not overlap.
This random selection of ten plots from within the main disturbed area
allowed for a representative initial characterization of the disturbed sur-
face as a whole, including impression types, dimensions, morphologies,
and density. Each plot was photographed along with individual imprints
of interest outside of the surveyed areas. Impressions were measured as
discreet, individual events. Impressions are superimposed and of such a
high density that it was not possible to assign individual depressions to
related groupings (trackways). All impressions within a plot were mea-
sured, inclusive of those only partially entering a given plot. Measure-
ments included impression length, width, and depth; the width and height
of surrounding sediment ridges; impression orientation; and such notable
features as shape, overprinting, and toe and claw prints.
Linear traces were photographed and measured for length, width, and
depth. Associated soft-sediment rims were measured for height and width,
orientation, and association with surrounding imprints. A 108-m strati-
graphic section was measured adjacent to the disturbed surface to record
eolian facies, sedimentary structures, bounding surfaces, and bioturbation
(Fig. 3). Data were analyzed for impression density, size distribution, and
predominant orientation. In all cases, specimens were viewed in situ and
samples were not collected.
RESULTS
Description of Disturbed Bedding Plane and Impressions
The Top Rock measured section (Fig. 3) provides stratigraphic context
for the disturbed surface. The section includes 51 cross-bedded cosets of
small (⬍3 m) to mega (⬎20 m) eolian dunes. Three distinct, heavily
bioturbated interdune deposits are located in interdune troughs between
larger dunes at 0–7 m, 61–66 m, and 107–108 m (Fig. 3). Within these
measured interdune intervals, intense bioturbation has obscured original
eolian stratification.
The focus of this study is an ⬃1-m-thick, massive, fine- to medium-
grained, white sandstone bed at 64 m in the measured section, in the
middle interdune deposit (Fig. 3). This interval is distinguished by its
proximity to a fluvial (wadi) channel, its high degree of bioturbation, the
presence of desiccation cracks, and the absence of cross-bedded, eolian
dune deposits. This interdune deposit is superimposed over the underlying
dune deposits such that the boundary is partially gradational. Impressions
penetrate an interdune surface that caps an eolian dune whose original
cross-bedding is nearly obliterated by bioturbation (Fig. 4). The density
of impressions can obscure the deflation plane. Horizontal and vertical,
unlined burrows increase upwards in this stratigraphic interval. Interdune
deposits up to 1 m thick occur above and adjacent to the imprinted sur-
face. These interdune bedding planes display nonbranching and branch-
ing, unlined, horizontal burrows lacking meniscate backfill, as well as
unlined vertical burrows and occasional desiccation cracks. A lens-
shaped, fluvial channel, ⬃4 m thick—with normally graded, fine- to me-
dium-grained sand, and bed thickness averaging between 0.15 m and 0.8
m—occurs laterally adjacent to the interdune deposit and stratigraphically
above the disturbed surface.
The Top Rock disturbed surface extends east–west over an area ⬃3000
m2 (⬃100 m long ⫻ ⬃30 m wide) and contains a high density of circular
to irregular impressions (Figs. 1, 5) numbering well into many hundreds.
PALAIOS
The densest concentration of impressions is in the easternmost 60 m (Fig.
1A). Penetration is commonly at an angle into the sediment (media), with
impressions ringed or surrounded by symmetric to asymmetric soft-
sediment ridges.
The survey records 473 impressions in 10 plots totaling 40 m2 with
an average density of 12 impressions/m2 (Table 1). Impressions have an
average length of 18.5 cm, with a range spanning from 1.5 to 140 cm.
In several instances, impressions larger than 100 cm are likely the com-
bination or merging of two imprints. Average impression depth is 4.9
cm, with a range between 0.5 cm and 30 cm. Impressions possess both
sloped and flat bases. Roughly one-third (152) of the impressions are
rimmed by lithified soft-sediment ridges of upturned sand (Figs. 1E, 5F,
H). Average ridge width is 4.2 cm (ranging between 1 and 18 cm); mean
ridge height is 2.2 cm (ranging between 0.5 and 11 cm). Fine-scale, soft-
sediment deformation is typically noted at the tip of asymmetric impres-
sions (Fig. 5A). Where impressions penetrate the media at an angle, ridg-
es are commonly best developed along the leading edge of penetration.
Seventy-one impressions provide a penetration orientation primarily to
the west-southwest (Fig. 6). Sixty-two impressions overprint atop another
impression.
Elongate, curvilinear traces up to 7.3 m in length and rimmed by par-
allel, soft-sediment rims were noted at several locations within the dis-
turbed surface. The impressions display a mildly sinuous course, with the
most easily distinguishable specimen (Fig. 1D, G) in the southwest extent
of the disturbed surface. These elongate impressions are 6 cm wide. Their
displaced, soft-sediment rims stand 10 cm wide by 3–4 cm tall.
INTERPRETATIONS
Interpretations of the deformed bedding plane are divided into three
categories and discussed in the order of: (1) individual impressions (trace
fossil footprints); (2) specific morphotypes and tail traces; and (3) the
collective interpretation of the numerous tracks as comprising a distinc-
tive trampled ground.
Track Occurrence and Preservation
The impressions are interpreted as vertebrate tracks based on several
important and distinguishing criteria that follow those of Martinell et al.
(2001):
1. Impressions are of the proper size range (tens of centimeters) for
vertebrate trackmakers.
2. Sixty-six (14%) of the impressions possess identifiable foot mor-
phologies. The remaining 86% of impressions are clearly similar and
related to those with identifiable foot morphologies. Repeated impression
morphologies are anomalous in a weathering or nonbiogenic regime, but
are consistent with a vertebrate origin (see discussion).
3. Impression floors are rarely flat and are commonly oriented at an
angle into the media, indicating a clear direction of travel to the west-
southwest (Fig. 6).
4. Thirteen percent of the impressions (62) display overprinting.
5. The impressions are constrained within one major bed (features are
temporally defined).
6. Other clearly documented Grallator, Brasilichnium, and Otozoum
dinosaur tracks on eolian slip faces are found in the same area (Loope,
2006).
7. Thirty-two percent of impressions (152) show symmetric to asym-
metric, primary plastic sediment deformation features (marginal ridges;
after Allen, 1997, and Gatesy, 2003), rimming impressions and at digit
tips (Fig. 5A, F, G, H). These features are penecontemporaneous with
impression formation.
A key to assigning a Jurassic, vertebrate origin to the Top Rock dis-
turbed surface is the soft-sediment marginal ridges related to the impres-
sions that indicate these are primary and syndepositional features, not
modern, late-stage, surficial weathering phenomena. The vertical relief
PALAIOS WET INTERDUNE DINOSAUR TRAMPLED SURFACE IN JURASSIC NAVAJO SANDSTONE
and asymmetric character of the ridges suggest they were formed by the
angular penetration of an object and the displacement-expulsion of ad-
jacent sand. Impressions lacking ridges are considered to be original, soft-
sediment features based on cross-cutting relationships and similarities be-
tween impressions with soft-sediment ridges and those without.
The abundance of marginal ridges and the preservation of small (⬍5
cm) prints along with larger prints indicate that many of the impressions
are true tracks as opposed to undertracks (e.g., Gatesy, 2003); the smaller
trackmaker would have insufficient mass to deform underlying beds. It
is possible that undertracks from previously overlying layers may imprint
the surface and account for depressions lacking definitive foot morphol-
ogy and marginal ridges. This combination of original tracks and under-
tracks may account for some of the variable quality in preserved tracks,
particularly in regards to the presence of pads, toes, claws, and digits.
Track Morphotypes
The Navajo Sandstone preserves a rich record of vertebrate footprints
but a paucity of body fossil remains (Irmis, 2005). Of the vertebrate tracks
in the Navajo Sandstone, those attributed to theropod dinosaurs are the
most common (Lockley and Hunt, 1995; Rainforth, 1997; Lockley et al.,
1998; Irmis, 2005). Additionally, traces assigned to a wide variety of
such vertebrate groups as Synapsida, Crocodylomorpha, Ornithischia,
Sauropodomorpha, and numerous other enigmatic footprints are cited as
preserved in this formation (Baird, 1980; Lockley and Hunt, 1995; Rain-
forth, 1997; Lockley et al., 1998; Carrano and Wilson, 2001; Yates, 2004;
Irmis, 2005). Although it is impossible to assign all impressions to ichn-
ogenera within the Top Rock disturbed surface—only 14% of tracks pos-
sess identifiable foot morphology—many can be assigned to the theropod
traces cf. Eubrontes, cf. Anchisauripus, and cf. Grallator, as well as that
of an unidentified sauropodomorph (J. I. Kirkland, personal communi-
cation, 2006; Fig. 5), based on size, morphology, and documented pres-
ence elsewhere in the Navajo Sandstone.
Grallator, Anchisauripus, and Eubrontes tracks are bipedal, tridactyl
impressions with less than 40⬚ divarication between digits II and IV
(Hitchcock, 1845, 1858; Rainforth, 1997; Olsen et al., 1998; Irmis, 2005).
Grallator impressions (Figs. 1C, 5B) are less than 18 cm long, Anchi-
sauripus are between 18 and 25 cm long, and Eubrontes tracks (Fig. 5D)
are classified when greater than 25 cm (Rainforth, 1997; Olsen et al.,
1998). Thirty-three tracks were assigned to these ichnospecies (Table 2).
In each case, digits are well defined, at times with pads and claw traces.
In this study, impressions are assigned to cf. Grallator, cf. Anchisauripus,
or cf. Eubrontes based on their known occurrence and documented char-
acteristics in Navajo Sandstone, tridactyl character, low divarication an-
gle, and dimension.
Another common morphology is an ovate impression with a clear, mid-
dle (III) claw deforming the sand (Fig. 5A, Table 2). This fine-scale
deformation at the apex of many ovate impressions is interpreted as the
tip of an elongated digit III belonging to a tridactyl trackmaker. Addi-
tional marks in the ovate track shaft correspond with digit II and IV of
a tridactyl footprint (Fig. 5A). Although the fine resolution of such foot
features as pads and toes may be obscured by other traces or poor pres-
ervation, the correlation of digit indentations on ovate shaft walls (Fig.
5A) with the occurrence of clear tridactyl footprints in the disturbed sur-
face (Figs. 1C, 5B, D) suggests that ovate tracks may also be related to
theropod footprints.
Tracks displaying manus and pes impressions (Fig. 5C, E, G) hold
closest affinity to sauropod tracks, but possess notable differences and
are, thus, assigned to a sauropodomorph. The impressions share the sau-
ropod characteristic U-shaped manus with a subcircular pes overprinted
atop the manus (Wilson and Carrano, 1999; Table 3). The interpreted
sauropodomorph tracks are quadrupedal with both manus and pes im-
pressions visible. Two basic morphologies can be described, one in which
the manus displays four toe or claw impressions arranged in a semicir-
cular arc (Fig. 5E), and the second in which individual toes or claws are
703
not clearly defined in the manus (Fig. 5C, G), but rather the manus is
crescent shaped and the pes is circular. These latter prints are remarkably
similar to compound structures described by Costa Da Silva et al. (2007,
fig. 4G) in the Caturrita Formation, Brazil, and attributed to prosauropods
similar to Pseudotetrasauropus (Costa Da Silva et al., 2007). Notably, in
both examples, the ovate pes trace diameter is less than the length of the
manus. The possible sauropodomorph impressions at the study site war-
rant further exploration and formal description with more extensive future
research.
Tail Traces
Elongate, curvilinear impressions extend up to 7.3 m; are rimmed along
their length by parallel, marginal soft-sediment ridges; and are interpreted
as protracted tail traces with expulsion rims (after Platt and Hasiotis,
2008). In areas of high impression-density, it was not possible to assign
individual, footprint-type impression associations to the tail traces. In an
area of lower track-density at the southwestern portion of the disturbed
surface, however, the association of the tail traces with dinosaur footprints
allows confident assignment of one trace to a quadrupedal dinosaur (Fig.
1D, G). This tail trace extends for ⬃4 m, is 6 cm wide, and 1.5 cm deep.
Soft-sediment expulsion rims are present along the impression’s length
on both sides. Following the classification established by Platt and Has-
iotis (2008), we interpret this trace as a protracted, simple tail trace dis-
playing high sinuosity. Associated quadruped footprints are circular, av-
eraging 7.7 cm in diameter, and comparatively shallow, only 1–2 cm
deep. Trackway width is ⬃0.4 m; stride length is ⬃0.5 m. As tail traces
are rarely preserved (Irby and Albright, 2002), their presence makes the
Top Rock surface unusual.
The Surface as a Trampled Ground
Five criteria are used to interpret the collective impressions of the Top
Rock surface.
1. The surface is similar to a Late Triassic dinoturbated interval de-
scribed by Costa Da Silva et al. (2007) in Brazil, particularly the striking
resemblance of Top Rock sauropodomorph tracks compared to compound
traces (two related impressions assigned to the same footprint).
2. Although claws and toes are only observed in 14% of the impres-
sions, marginal ridges in 32% of impressions are consistent with models
of Allen (1997), Gatesy (2003), and Henderson (2006).
3. Although the Top Rock surface formed in fine- to medium-grained
sand as opposed to muddy sands to clays, the disturbed surface shows
noticeable similarity to trampling described by Laporte and Behrensmeyer
(1980) and Ashley and Liutkus (2002). Trampling described in plan view
by Laporte and Behrensmeyer (1980) is characterized by impressions up
to 15 cm deep, depression floors tilted in the direction of animal move-
ment, raised sediment rims surrounding imprints, poor preservation of
well-defined footprint morphology, and the ability to identify only single
footprints and short trackways.
4. The impressions are of such a high density that the large number
of prints obscures identification of individual trackways.
5. Although the overall preservation of footprints in wet sands differs
from preservation in cohesive muddy sand and clay-dominated soils, the
Top Rock interdune area provides ideal conditions for the preservation
of a vertebrate trampled ground. Moist, possibly vegetation-free sand (La-
porte and Behrensmeyer, 1980; Costa Da Silva et al., 2007), and the
potential for a rapid increase in sedimentation provided by eolian dune
shifting create favorable preservation conditions (Ashley and Liutkus,
2002). The disturbed bedding plane surface with numerous circular to
ovate, high-density, soft-sediment impressions with distinct to modified
footprint features is, thus, interpreted as a trampled surface produced by
numerous footprint impressions made while the sediment was soft. Con-
ditions of moist sand within an interdune setting contributed to the pres-
ervation of this trampled surface.
704 SEILER AND CHAN PALAIOS

FIGURE 2—Ten individual measured plots to characterize the trampled surface delineated by dashed line (2 m/side). Scale bar ⫽ 1 m. A) Plot 1. B) Plot 2. C) Plot 3.
D) Plot 4. E) Plot 5. F) Plot 6. G) Plot 7. H) Plot 8. I) Plot 9. J) Plot 10. See Table 1 for measurement data.

The trampled surface displays a moderate degree of dinoturbation not be individual footprints—some may be partial toe imprints or sedi-
based on the scale of Lockley and Conrad (1989). The high density and ment splatters associated with foot removal from the sand media. Cross-
abundance of impressions of various sizes (from 2 to ⬎50 cm) allows cutting relationships, however, imply that all impressions are related to
for the likelihood that all impressions, particularly the smaller ones, may the same (temporal) track-forming episode.
PALAIOS WET INTERDUNE DINOSAUR TRAMPLED SURFACE IN JURASSIC NAVAJO SANDSTONE
FIGURE 3—Stratigraphic column including the Top Rock trampled surface. The
trampled surface is in the middle interdune deposit with a high degree of bioturbation
surrounded by medium-sized dune sets. Grain size varies from fine- to coarse-grained
sandstone.
A Vertebrate Origin versus Alternate Interpretations
In examining the interpretation of the disturbed bedding surface as a
trampled bed, alternate interpretations were considered. Three categories
of alternate interpretations are presented: (1) weathering features, (2) non-
biogenic soft-sediment deformation, and (3) other biogenic related struc-
tures. Each alternative is described and shown to be inconsistent with the
features observed at Top Rock.
Potholes, or weathering pits, are relatively recent surficial features gen-
erally occurring as broad, shallow, flat-bottomed depressions on flat to
gently sloping outcrops (Netoff et al., 1995). Chan et al. (2005) identified
pothole occurrence along three genetic surfaces: (1) at the exposed top
of a formation; (2) on exposed regional supersurfaces (Kocurek, 1996);
and (3) on exposed, first-order, eolian bounding surfaces where interdune
deposits are common. Netoff et al. (1995) identified physical, chemical,
and biological forces that account for pothole formation. These include
water- and wind-dependent weathering, granular disintegration induced
by mineral crystallization, clay desiccation and hydration, freeze and thaw
705
FIGURE 4—Schematic cross-section of the Top Rock trampled surface. Impressions
penetrate the wetted interdune deflation surface of a heavily burrowed eolian dune.
Interdune deposits directly overlie the trampled deflation surface. Bioturbation in-
creases upward in the heavily burrowed dune facies.
cycles, carbonation, dissolution, hydrolysis, and biological activity. Pot-
holes and pits have gently sloping borders when located on open surfaces,
where bedding lacks joints and regular water flow. By any account, pot-
holes are weathering features that postdate deposition and develop after
sediment has been lithified.
The track impressions at Top Rock are inconsistent with weathering
pits and potholes in several regards:
1. Impression floors are rarely flat and are typically oriented at an
angle into the sandstone.
2. The tracks are steep walled but lack the hydraulic mechanism or
abrading tool required for carving steep-sided potholes, and no pebbles
occur within impressions
3. The impressions lack a modern fluvial or even small rivulet erosive
force common to many potholes formed in both sandstone and igneous
rocks and do not display the smoothed chutes and pools common in the
surficial, water-scoured Navajo Sandstone. Likewise, the impressions lack
the geomorphic character (steps and risers) common of eolian weathering
elsewhere at Coyote Buttes (Loope et al., 2008).
4. Well-formed lichens on impression walls suggest the features are
not being actively scoured by wind or water.
5. Imprint population density far exceeds that of potholes observed on
the Colorado Plateau. Likewise, impressions are superimposed and do not
display the distribution common of weathering pits (Chan et al., 2005).
6. Repeated impression morphologies occur that would be anomalous
in a weathering regime, but consistent with a vertebrate origin.
7. Finally, the lack of high-density potholes on any other surface in
the area and the prevalent soft-sediment ridges around the impressions
indicate these cannot be original weathering potholes, although some wa-
ter collection and black biofilms are superimposed on the original, pri-
mary depression structures interpreted as footprints.
Other such potential weathering phenomena as taphoni—honeycomb
weathering structures—generally occur at a smaller scale than the Top
Rock impressions and are commonly restricted to vertical faces. Taphoni
structures require salt, biologic crusts, and porosity differences in the
sandstone (Turkington 1998; Campbell, 1999; Nash, 2000) to facilitate
706 SEILER AND CHAN PALAIOS

FIGURE 5—Representative tracks and impressions—measured format is length ⫻ width ⫻ depth (where known). Insets contain diagrams of interpreted features. A) Ovate
print 42 ⫻ 29 ⫻ 11.5 cm. Marginal ridge 3 cm wide ⫻ 2 cm tall. Digit III deformation (arrow) 8 cm long. Locations of inferred digit II and IV markings indicated by
white triangles. B) cf. Grallator track 11 ⫻ 5.5 ⫻ 2.5 cm. C) Possible sauropodomorph track 18 ⫻ 20.5 ⫻ 12 cm. Note preservation of fine-scale soft-sediment deformation
at distal edges of manus. D) cf. Eubrontes print ⬃42 ⫻ ⬃38 cm. M ⫽ metapodium. E) Sauropodomorph track 36 ⫻ 44 cm. The ovate pes is 23 cm ⫻ 20 cm and is
imprinted significantly deeper than manus. F) Tracks displaying asymmetric soft-sediment marginal ridges. Inset diagrams the soft sediment ridge configuration. G) Sau-
ropodomorph, 15 ⫻ 18 cm displaying manus, pes, and asymmetric marginal ridge. H) Well-developed primary structures that distinguish the tracks. SW ⫽ steep walled;
AP ⫽ angular penetration; MR ⫽ marginal ridge; SSD ⫽ soft-sediment deformation at digit tip; arrows point to possible toe impressions. I) Rivulet channel (arrow) drains
from higher (top) to lower (bottom) impression.

weathering. Porosity within the trampled surface is relatively constant,
with no significant salt source present in the area. Taphoni is not recog-
nized elsewhere in the field area, making this an unlikely explanation for
the impressions.
Possible synsedimentary processes that could lead to features similar
to the dinosaur tracks include nonbiogenic soft-sediment deformation.
The disturbed surface, however, displays neither the vertical pillar struc-
tures expected from a synsedimentary, dewatering origin nor the irregular
convolutions common of soft-sediment deformation events when viewed
in plan view. Bedding-scale disruptions are not traceable lateral to the
trampled surface, as would be expected from large-scale deformation
events (e.g., ground shaking). Preserved burrow structures and faint eo-
lian laminae packages near the impressions within the trampled surface
also indicate a lack of major, large-scale dewatering or ground-shaking
disturbances.
Additional alternate interpretations of biogenic origins for the disturbed
bed could include other vertebrate or invertebrate burrowing, casts around
preexisting biogenic structures, fish nests, and ray traces. Invertebrate
burrow structures commonly display burrow lining, branching or bifur-
cations, or tiering (Bromley, 1996). Large vertebrate burrows from the
Middle Jurassic Entrada Sandstone described by Loope (2006) occur at
a lower angle (16⬚–22⬚ from horizontal) than the impressions and display
PALAIOS WET INTERDUNE DINOSAUR TRAMPLED SURFACE IN JURASSIC NAVAJO SANDSTONE 707

TABLE 1—Trampled surface data from ten plots. Plots correspond to areas shown in Figure 2.

Plot 1 Plot 2 Plot 3 Plot 4 Plot 5 Plot 6 Plot 7 Plot 8 Plot 9 Plot 10 Total

Impressions 43 34 67 50 80 20 61 34 66 18 473
Circular 38 21 38 18 13 1 11 6 10 2 158
Ovate 0 4 10 10 29 13 33 20 33 12 164
Elongate 0 0 0 3 17 2 4 3 9 1 39
Irregular 5 9 19 19 21 4 13 4 13 1 108
Tails 0 0 0 0 0 0 0 1 1 2 4
Mounds 6 14 20 20 29 5 19 12 22 5 152
Overprints 23 0 5 3 15 1 7 2 5 1 62
Oriented 8 7 9 7 10 7 8 3 10 2 71
Tridactyl 5 2 3 4 5 1 6 2 5 0 33
Ovate with digit III 0 3 4 0 5 1 3 2 4 4 26
Sauropodomorph 0 0 2 2 2 0 1 0 0 0 7
Total with definitive print morphology 5 5 9 6 12 2 10 4 9 4 66

burrow fill not observed at the trampled surface. If the impressions were
burrows, where overlapping occurs, one would expect to see collapse of
the preceding burrow into the new, crosscutting example; this is not ob-
served at the trampled surface. Similarly, burrows do not incorporate
adjoining sediment into marginal ridges (Lea, 1996) observed around
vertebrate footprints. The large impressions of Top Rock are of an in-
compatible size to typical small invertebrate and vertebrate burrows from
eolian settings (Albrandt et al., 1978; Ekdale and Picard, 1985; Ekdale
et al., 2007).
Parrish et al. (2007) document coniferous trees preserved from nu-
merous Navajo Sandstone interdune deposits with some casts that can
form in sandstones. The criteria of Costa Da Silva et al. (2007) used to
interpret similar Brazilian impressions as dinosaur traces can be applied
here: root and branch structures are missing and internal woody structure
is lacking, as are bark impressions and films. Although fish nests and ray
traces can form subaqueous impressions in standing water, fish and ray
traces are not consistent with the eolian interdune setting. Furthermore,
their individual impressions would not overlap or display marginal rims
(Feibel, 1987; Martinell et al., 2001).
None of the alternate interpretations considered here are consistent with
the observed field data in the study site. The strength of the interpretation
of dinosaur trackmakers for a trampled bed in an eolian interdune setting
lies in the clearly preserved soft-sediment impressions consistent with
vertebrate footprint size and morphologies, as well as associations with
other documented scattered dinosaur tracks in the area.
DISCUSSION
Track Size Distribution
Tracks with identifiable foot morphologies at the Top Rock study site
do not yield clear and distinct size groupings, but rather show a grada-
tional transition from cf. Grallator to cf. Anchisauripus to cf. Eubrontes
(Table 2). This can be attributed to five factors: (1) the presence of species
of roughly overlapping size—distinct foot impressions imply the exis-
tence of at least four different species; (2) the presence of juvenile to
adult individuals of a particular species; (3) preservation method; (4)
partial footprint impression; and (5) a combination of the above. Regard-
less, numerous track types and print sizes in the interdune indicate that
a diversity of taxa was supported within the Navajo erg.
There is considerable discussion over whether tracks of a particular
size can be generalized to a certain depositional facies—small tracks con-
strained to cross-bedded dune foresets, large tracks constrained to inter-
dune deposits—or whether such occurrence is merely a bias in preser-
vation (Rainforth, 1997; Irmis, 2005). In this case, tracks of both large
and small fauna are preserved in the interdune setting. The abundance of
both large and small tracks indicates that, at the least, the habitat of small
track-bearing organisms (i.e., Grallator trackmakers) extended from
cross-bedded dune sands, into an interdune setting occupied with large
track-producing animals. The Top Rock trampled surface, thus, supports
the idea that correlating track sizes to eolian facies reflects a bias in
preservation.

FIGURE 6—Rose diagram showing dominant west-southwest direction of travel
perpendicular to northwesterly paleowind.
Paleobiologic Implications
Winkler et al. (1991) suggested that the existence of both body and
trace fossils at several interdune deposits in the Navajo Sandstone sup-
ports the presence of a well-developed food chain requiring significant
time for establishment. The food web implied by the abundant and varied
interdune track types at Top Rock reflects the long-lived nature of inter-
dunes in the Navajo erg. The resources required to support a complex
food web of large organisms implies that interdunes did not exist in
ecologic isolation of one another. Animals may have traveled significant
distances through the erg between interdunes.
The orientation of tracks (Fig. 6) to the west-southwest (240⬚–250⬚)
lies perpendicular to the predominant northwest paleowind (Parrish and
Petersen, 1988; Chan and Archer, 2000; Loope et al., 2001) and parallel
to dune crests (striking 39⬚). This implies that travel through the trampled
surface was directed by the orientation of surrounding dunes. Subordinate
orientations may indicate wandering within the interdune area or behav-
ioral dynamics.
708 SEILER AND CHAN PALAIOS

TABLE 2—Tridactyl and ovate impressions identified from plots in Table 1. All measurements in cm.

Ichnospecies Length Width Depth Mound width Mound height

Clear tridactyl impressions

cf., Grallator Average 10.3 8.8 3.0 3.4 1.6
n ⫽ 20 Range 5 to 17 4.5 to 17 1 to 10 1.5 to 9 0.5 to 4
cf., Anchisauripus Average 21.0 18.3 6.5 1.5 1.0
n⫽3 Range 20 to 23 15 to 24 3.5 to 11 1.5 1.0
cf., Eubrontes Average 41.6 34.9 7.5 3.3 2.6
n ⫽ 10 Range 25 to 78 10 to 78 3 to 15 1.5 to 5 1 to 6
Ovate impressions with clear, middle (III) claw impressions
cf., Grallator Average 13.0 8.8 6.4 1.8 1.0
n⫽6 Range 6 to 17 6 to 13 4 to 10.5 1 to 2.5 0.5 to 1.5
cf., Anchisauripus Average 21.5 17.1 6.5 4.4 2.2
n⫽7 Range 19 to 23 14 to 23 5 to 8 2.5 to 9 1 to 3
cf., Eubrontes Average 35.9 24.3 10.0 4.6 1.7
n ⫽ 13 Range 25 to 52 18 to 39 6 to 20 2.5 to 8 1 to 2.5

Finally, dinosaur tracks in the Navajo Sandstone are significantly more
abundant than body fossils, which are preserved rarely (Irmis, 2005; D.D.
Gillette, personal communication, 2007). Gillette (personal communica-
tion, 2007) further asserts that little is known of dinosaurs in North Amer-
ica until later in the Mesozoic when a diversity of megafauna suddenly
appears in the Late Jurassic Morrison Formation (Winkler et al., 1991).
The discovery of the Top Rock trampled surface suggests that species
distribution and dynamics can be extended deeper into the Early Jurassic.
Paleoclimatic Implications
Although the Navajo Sandstone records predominantly arid conditions,
the well-defined angular penetrations and soft-sediment deformation ridg-
es of the Top Rock trampled surface (Figs. 1E, 5F–H) argue for the
existence of a wet interdune paleoenvironment. Any tracks penetrating
dry interdune sands would have collapsed upon foot extraction. It is rea-
sonable to expect that the reworking of dry sand by wind would erase
the majority of any dry marginal ridges surrounding print impressions.
The deposits of three, long-lived pluvial periods are described at the
Coyote Buttes (Loope and Rowe 2003; Loope 2006). The lowermost
interdune deposit measured in this study (0–7 m, Fig. 3), likely corre-
sponds to pluvial episode 2 of Loope and Rowe (2003). The middle
interdune deposit (61–66 m), in which the trampled surface is located,
likely correlates to pluvial episode 3 (Loope, 2006, fig. 1). A final, small
exposure at the top of the measured section (107–108 m) may signal a
fourth pluvial episode. Pluvial episodes associated with interdune deposits
indicate large climatic shifts were present during the Jurassic in the Na-
vajo erg—a time when continental and marine sedimentary and fossil
data suggest that there was a uniform climate throughout the continents
(Schaeffer, 1970; Hallam, 1993).
A change to a wetter climate may have allowed water table recharge
to interdunes (Loope and Rowe, 2003). Given the high porosity of eolian
sand, it is also reasonable to expect that precipitation into an interdune
would quickly infiltrate unless the water table was present at or above
the ground surface. In one track, a small trough leading to a lower im-
pression (Fig. 5I) indicates the active draining of water from the higher
print—possibly as the animal’s weight exuded interstitial water from the
sand. Variations in impression depth may also be linked to sediment water
content and track density. Where track density is greater, impressions are
deeper, possibly due to the progressive liquefaction of the sediment with
increased trampling. Following Lockley’s (1986) assertion that a direct
relationship exists between track depth and sediment water content for
animals of a given weight in a homogeneous medium (provided the clean
quartz arenite), it is reasonable to expect that future modeling may yield
accurate estimations of both the animal’s size and weight and the inter-
dune water content. This relationship may be elaborated upon further
through the examination of the soft-sediment marginal ridges surrounding
impressions.
Dinoturbation as a Mechanism for Convolute Bedding
Although Navajo Sandstone at the Coyote Buttes displays many areas
of convolute bedding, in places exceeding 10 m thick, the trampled sur-
face offers an explanation for bedding convolution at the smaller, sub-
meter to meter scale. Convolute bedding and soft-sediment deformation
in the Navajo Sandstone can be extensive and has been attributed to
factors including earthquake liquefaction, a fluctuating groundwater table,
gravity flow, and current shear (e.g., Doe and Dott, 1980; Horowitz, 1982;
Bryant, 2003; Loope and Rowe, 2003; Bryant et al., 2006). Bedding
convolutions caused by dinosaur tracks would be more localized and at
a smaller, meter scale. Although individually defined tracks are not visible

TABLE 3—Sauropodomorph impressions. Note that three distinct, well-preserved sauropodomorph impressions (2 small, 1 large) that occur adjacent to, but outside of measured
plots, were included in this characterization. All measurements in cm.

Pes length Pes width Pes depth Manus length Manus width Manus depth

Small sauropodomorph impressions n ⫽ 6

Average 14.3 12.8 8.0 20.9 3.7 4.9
Range 12 to 16 10 to 15 4 to 12 20 to 24 3 to 5 2 to 12
Large sauropodomorph impressions n ⫽ 4
Average 24.3 18.5 5.5 33.5 7.5 3.3
Range 23 to 27 14 to 23 4 to 6.5 28 to 44 5 to 10 2.75 to 4
Overall sauropodomorph impressions n ⫽ 10
Average 19.3 15.6 6.5 26.0 5.2 4.2
Range 12 to 27 10 to 23 4 to 12 20 to 44 3 to 10 2 to 12
PALAIOS WET INTERDUNE DINOSAUR TRAMPLED SURFACE IN JURASSIC NAVAJO SANDSTONE
in an oblique view of smaller deformed beds at the Coyote Buttes, the
density, size, and spacing of convolutions approximates that of trampled
surface tracks. Provided that dinosaur tracks are a possible cause for
small-scale convolutions, the track record in the Navajo Sandstone may
be much more numerous than previously acknowledged. Linking submeter-
to meter-scale convolutions to biologic trackmakers significantly increases
the potential for identifying large megafauna traces and migration pathways
during the Early Jurassic in the Navajo Sandstone.
This initial survey documents the presence of a unique, wet interdune
trampled surface. Future aerial mapping and modeling analyses could also
yield accurate estimations of animal size and behavior while assisting in
distinguishing individual trackways amongst the high concentration of
prints. The trampled surface, thus, is a valuable tool for elucidating de-
tailed paleobiologic, paleohydrologic, and paleoclimatic proxies for the
Jurassic of the Western Interior.
CONCLUSIONS
An unusual ⬃1-m-thick surface showing extensive (⬎1000), high-
density, steep-walled impressions from soft-sediment deformation occurs
in an interdune interval of the Jurassic Navajo Sandstone at the Coyote
Buttes along the Arizona-Utah border. This distinctive bed preserves a
trampled surface created by dinosaurs that visited the wetted eolian site.
The repeated foot morphologies, evidence of overprinting, angular pen-
etration, west-southwest alignment, abundance of soft sediment-
deformation features—marginal ridges and deformation at digit tips—and
impression density argue convincingly for a dinosaur origin of the sur-
face. This dinosaur-trampled surface at Coyote Buttes is unusual in ex-
hibiting a combination of multiple overlapping track types and sizes,
footprint features including claws and toes, high track density, and rare
tail traces. Patterns of at least three ichnogenera are present including cf.
Eubrontes, cf. Anchisauripus, cf. Grallator, and the tracks of a sauro-
podomorph.
The well-preserved tracks of the Top Rock trampled surface provide
several important paleobiology, paleohydrology, and paleoclimate prox-
ies: (1) dinoturbation is a mechanism for producing localized submeter-
to meter-scale convolute bedding in moist eolian sediment; (2) the south-
western margin of this Jurassic paleodesert provided sufficient resources
to support a diverse population of megafauna; and (3) the trampled sur-
face provides additional evidence of a pluvial climate episode, during
which shallow groundwater flowed into interdune areas.
The Coyote Buttes area holds significant scientific, educational, and
aesthetic value for resource management. The remarkable cyclic eolian
stratification accentuated by vivid diagenetic coloration, a gracefully
scoured landscape, and the record of important dinosaur occupation dis-
tinguishes the Coyote Buttes from many other Navajo Sandstone locales
across the Colorado Plateau.
ACKNOWLEDGMENTS
We gratefully acknowledge John R. Bowman, William T. Parry, Erich
U. Petersen, Anthony A. Ekdale, Bucky Gates, Lindsay Zanno, Jessica
Allen, and Sally Potter at the University of Utah, as well as Jim Kirkland
at the Utah Geological Survey for their input to this study. The Vermilion
Cliffs National Monument administration, as well as Rody Cox and Mike
Salamacha at the Bureau of Land Management, provided valuable logis-
tical and permitting assistance (Permit RP07VCNM06). Thanks to Brian
Parnham and Sara Hanner for help in the field and to David Gillette, Loı̈c
Costeur, and anonymous reviewers for critical review of the manuscript.
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ACCEPTED MAY 27, 2008