Recognizing and Utilizing Vertebrate Tracks in Cross Section: Cenozoic Hoofprints from

Nebraska
Author(s): David B. Loope
Source: PALAIOS, Vol. 1, No. 2 (Apr., 1986), pp. 141-151
Published by: SEPM Society for Sedimentary Geology
Stable URL: http://www.jstor.org/stable/3514507
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 REPORTS
RESEARCH
REPORTS
RESEARCH
Recognizing and Utilizing
Cross
Cenozoic Hoofprints from
DAVIDB. LOOPE
Departmentof Geology,Universityof Nebraska,Lincoln,NE 68588-0340
PALAIOS,1986, V. 1, p. 141-151
Because vertebratetracksexposedin verticaloutcropsare fre-
quentlyoverlookedor misinterpreted as physicallyinduceddefor-
mation structures,importantpaleoecologicand sedimentologic
informationremainsuntapped.Laminationswithineoliandune
sands of the NebraskaSand Hills (Holocene)and ephemeral-
streamdepositsof the lowerArikareeGroup(late Oligocene)are
commonlybrokenor sharplydownwarpedto form isolated or
paired, concave-upstructuresthat varyfrom 4 to 22 cm in
diameter.A central ridge divides the lower portion of some
structuresinto two distinctlobes.Althoughbedding-plane expo-
sures are rare in bothdeposits,extensivesearch revealedsome
structuresin linear alignment,confirminga biogenicoriginfor
the deformation.TheHolocenetracks,probablymade by bison,
wererepeatedly producedduringthemigrationof thelargeeolian
bedforms,suggesting that food and water were available in
interduneareas. Smectitegrain coatings made surface sands
cohesive,therebystronglyinfluencingtrackmorphology and en-
hancingpreservationpotential.Oligocenetrackswereproduced
in veryfine sand by severalspeciesof hoofedvertebrates.Close
verticalspacingof track-bearingbedsrevealsthatmostsediment
accumulatedin relativelythinpackages.Thegeneralabsenceof
heavilytrampledhorizons(as wouldbe expectedalong diastems)
may be the result of rapid consolidationof sands by evaporite
cementation.Tracksmayhavebeenproducedduringbriefinter-
vals of time immediatelyfollowing depositionand preceding
cementation.
INTRODUCTION
Vertebratetrackwaysare a well-knownsource of paleonto-
logic andsedimentologicinformation(Sarjeant,1975). Perhaps
becausevertebratetracksare so easy to recognizeon bedding-
planeexposures, untilrecentlylittleattentionhas been givento
theirappearancein verticalsection. Photographsanddrawings
publishedby Vander LingenandAndrews(1969), McKee and
Bigarella(1972), Lewis and Titheridge(1978), Laury(1980),
and Hunter et al. (1984) have, however, documented the
Copyright? 1986, The Society of EconomicPaleontologistsand Mineralogists
141
141
Vertebrate Tracks in
Section:
Nebraska
deformationthat takes place when large animalsmove across
soft, laminatedsediment. On the basis of their observationsof
Quaternarydepositsin East Africa,LaporteandBehrensmeyer
(1980) have recently argued that large vertebrates have the
potentialto rework terrestrialsediments to the same extent
that benthicinvertebratesreworkmarinestrata. Preservation
of tracksrequirescompactiblesubstratesthat are accessibleto
vertebrates; rates of tramplingand burial control whether
sediments record individualtracks or are totally bioturbated
(Laporteand Behrensmeyer, 1980, fig. 4a).
The Holocenetracks describedhere are from exposures of
duneandinterdunedepositswithinthe NebraskaSandHillsand
were probablyproducedby bison. Oligocene hoofprintsare
well exposed in outcropsof fluvialsheet-flooddeposits within
the GeringFormation(ArikareeGroup)at Scotts BluffNational
Monumentin westernmostNebraska;these trackswere made
by several differentspecies of cloven-hoofedmammals.The
morphologyof both Holocene and Oligocene hoofprintsindi-
cates that they were produced in cohesive sand. Much of
surfacesand in the SandHillsis cohesive even when dry, due
to clay coatingson grains;such coatingsgreatlyenhancedthe
preservationpotentialof the buriedtracks. Individualtracksor
pairs of tracks in the Holocene and Oligocenesediments are
typicallywidely spaced laterally,but closely spaced vertically.
The verticalspacingof track-bearinglaminaeallowsdivisionof
stratainto discrete depositionalpackages. Close verticalspac-
ing suggests that sand-drivingwinds deposited relativelythin
sedimentpackagesin the NebraskaSandHillsand that tracks
were producedby resident, ratherthanmigratorypopulations.
Much of the Gering Formationis composed of similarlythin
packagesthataccumulatedwithinan ephemeralstreamsystem
dominatedby sheet-floodevents. The lackof heavilytrampled
zones (whichmight be expected to mark diastems) suggests
that the Holocenesedimentaccumulatedsteadily;in contrast,
the same patternin the Oligocenerocks couldbe the result of
rapidcementationof newly depositedsandby evaporites.The
abundanceof vertebrate tracks in these strata shows that,
under certain conditions, eolian dune fields and ephemeral-
streamfloodplainscan be very favorablesites for preservation
of vertebratetracks.
0883-1351/86/0001-0141/$03.00
142 LOOPE

430
ILc
r-

--- cc"l` ..-

~L- 'd r

41?

?rrS;r*e .:~ :" I----- 7- 4~,-.----~;

102?

FIGURE1-Map showing outline of the Nebraska Sand Hills and L

I .
location of Burwell (B), Thedford (T), Valentine (V), and Scotts,Bluff
J,
National Monument (SB).

The mainpurposes of this paper are: 1) to documentsedi- FIGURE

2-Concave-up deformation structures in large-scale cross-
mentarystructuresproducedby the hooves of large mammals stratified Holocene dune deposits at Calamus River dam site near
in nonmarinesands and sandstones of central and western Burwell, Nebraska. Note that deformation occurs throughout vertical
Nebraska;2) to providecriteriaby whichthese featurescan be extent of exposure.
differentiatedfromnonbiogenicdeformationstructures;and3)
to further demonstrate the utility of vertebrate tracks for
sedimentologicaland paleoecologicalinterpretations. winds swept ripples across surfaces with dips well below the
angle of repose.
HOLOCENEOF THE NEBRASKASAND HILLS
Descriptionof Tracks
Locationand GeologicSetting The features interpretedhere as vertebrate tracks are con-
cave-up deformationstructuresthat are circularto oval in plan
Occupyingan area of 57,000 square kilometers, the Ne- and
braska Sand Hills (Fig. 1) are the in range from7 to 16 cm in diameter(Figs. 3, 4, 5). Near the
largest dune field the
Western Hemisphere (Smith, 1965). This dune field, now tops of structures, laminaeare abruptlytruncatedor sharply
stabilizedby prairievegetation, is composed of simple and folded(Figs. 3, 4a). As in the structuresreportedby Van der
compound,transverse to obliquebedformsthat reach heights ward.LingenandAndrews(1969), concavitydies out graduallydown-
up to 100 m (Ahlbrandt andFryberger,1980). Carbon-14dates Althoughdeformationextends verticallyas much as 25
cm, individuallaminaeare displacedno more than15 cm. Within
reported by Ahlbrandtet al. (1983) from sediments directly some
beneath the dunes suggest that the dune field is primarily individualstructures, a central ridge divides the lower
Holocenein age. The sedimentarystructureswithinthe dunes portioninto two distinctlobes, whichare visible in both cross-
are visiblein a largenumberof exposures throughoutthe Sand sectionaland planviews (Fig. 4). The interiorsof many struc-
Hills (Ahlbrandtand Fryberger, 1980). Most of the observa-
tions for this paperwere made near the eastern marginof the
Sand Hills at excavations for the CalamusRiver dam near
Burwell, Nebraska, and at blowouts and stream cuts near
Thedfordin the central Sand Hills (Fig. 1). Structuresinter-
preted here as vertebrate tracks are present throughoutthe
Sand Hills; many are recognizablein publishedphotographs
from the northern and central Sand Hills (Ahlbrandtand
Fryberger, 1980, figs. 9, 10, 11, and 13). At all vertically
extensive exposures, it is clear that the tracks are not re-
strictedto the upperportionsof the dunes, but are distributed
throughoutthe thicknessof the eoliandeposit (Fig. 2).
All deformationstructuresinterpretedas hoofprintsare de-
velopedin thin,inverselygradedlaminaedippingbetween 0 and
24 degrees. These laminaewere clearlydepositedby migrating
windripplesand, in the terminologyof Hunter(1977), can be
classifiedas subcriticallyclimbingtranslatentstrata. Laminae
with very low dips accumulatedin interduneareas or as sand
sheets (Frybergeret al., 1979); more steeply dippingstrata FIGURE3-Tracks in horizontal wind-ripple laminae. Interdune or
were deposited on leeward slopes of bedforms, where side sand-sheet deposit near Burwell. Pen is 13 cm in length.
 .

HOOFPRINTS
CENOZOIC 143

N=41

LL

I
2 4 6 8 10 12 14 16

TRACK DIAMETER(CENTIMETERS)

j"~n
iN!
,,Hs*'" FIGURE Size-frequency data for Holocene tracks measured from
AFIGURE......~~~~verti cal exposures, NebraskaSand Hills. Some variationin diameter
AtI"
:Ar
....-:e?o: ~ .. .. is result of oval plan of structures(Fig. 4b).

~ '
l'-i:;:':"!",.- -~,! b,,."/.the Sand Hills, fortuitous exposures and small excavations
providedglimpses of short trackways(Fig. 6) that along with
the blgobedmorphologyof some structures, confirmedthe
.." . ' biogenic hypothesis. After the structures are recognized as
,:
tracks, the distributionof individualdeformationstructuresand
other aspects of their morphologyprovideimportantsedimen-
tologicaland paleoecologicalinsights.
ContrastwithNonbiogenicStructures
The size-frequencydistributionof the deformationstructures
(Fig. 5) stronglysupportsthe biogenicinterpretation:none of
appearing32the structuresare too large or too small to be tracks. When
closely spaced, however, the SandHillsstructuressuperficially
resemble convolute bedding, a structure that Allen (1982)
defines as a "lateray extensive series of more or less regular
exterally
folds developedthroughoutor confinedto the upperpart of a
single sedimentationunit." Convolute bedding, however,i is
~with
4 (d-A)17FIGo~~~~uRE
sFIGURE ~leTrack
dee wly
4-AuTac
wthvetiypical vertical
wallstrdandbilobedstlaower
portion.o
Ir due c B dati ENo
th sedimntatonuit.etres to root
rapid
sedimentation,
Surfaceconcavitywas infilledby coarse-grained lag deposit.Note typicaly developed in strata that, due to rapidsedientation
raised "rim"at upper margins. Subhorizontallines crossing struc- were originally very loosely packed (Allen, 1982, p. 343).
tures are post-depositionalaccumulationsof silt and clay (soil lame- Accordingly, this structure has been used as evidence for rapid
Ilae or "dissipationstructures"of Ahlbrandtand Fryberger,1980). B) deposition (Collinson and Thompson, 1982, p. 145). Studies of
Plan view of bilobed track near the middle of a three-meter-thick
crossbed set. ?'*' : . '
' , R,
_ '.i:

tures are composedof materialthat is texturallydistinctfrom t ._ .

the underlyingand surroundingdeformed sediment. Com-
monly, this central portion contains very coarse sand and- ' -

granules(Fig. 4a); in rare cases, a thin (about5 mm) layer of .:! -* ... S
":
silt is preserved on the floor of the concave-upstructure.
Along individualstratigraphiclevels, concave-upstructures _, -
are laterallyisolatedor in pairs. The structuresare commonly
closely spacedvertically(Fig. 3) andmay dominatethe aspect -"
'
of outcrops exposing several meters of strata (Fig. 2); laterally ;' -
-
'
adjacentbeds may lack such deformation. F

Interpretation
In contrast to the easily recognized vertebrate trackways
appearingon bedding planes (Sarjeant, 1975), most tracks
appearingin cross sectionhave probablybeen eitherignoredor FIGURE
6-Part of shorttrackway
in large-scalecross-stratified
sand,
misinterpreted (Lewis and Titheridge, 1978). Tracks in vertical Calamusdam site, Burwell,NE. Note lack of root traces in laminae
outcropscan closely resemble other types of deformation.In belowtracks.Machetehandleis 15 cm long.

I
144 LOOPE

moder wind-rippledeposits indicatethat the stratacontaining

the SandHillsdeformationstructureshad low initialporosities
and were deposited relatively slowly. Because of strong
differencesin grainpackingand porosity, the differenteolian
stratificationtypes vary in their susceptibilityto deformation
(Bagnold,1941; Hunter, 1981). Avalancheor grainflowstrata
arelooselypacked;the large-scaledeformationcommonlyfound
in ancient eolian sandstones is commonly concentrated in
deposits of this type, because their high porosity allows
liquefaction.In contrast, strata dominatedby climbing-ripple
deposits-apparently due to their low initialporosities-are
rarelyinvolvedinthiskindof deformation(Doe andDott, 1980).
In lightof the evidence frommodem andancienteoliansands,
the abundanceof relativelylarge-scaledeformationstructures
in the wind-rippledeposits of the Sand Hills initiallyseemed
incongruous.It was this paradoxthatled to the hypothesisthat
the deformationstructureswithinthe typicallystablestrataare
biogenic.
AhlbrandtandFryberger(1980) explainedsome deformation
structures within eolian strata of the Sand Hills (which are
identicalto the ones interpretedhere as tracks)as the result of
compressionat the base of slipfacedeposits. Several workers
have reportedobservationsof this type of deformation.From
the Coorongregion of southernAustralia,Brown (1969) de-
scribedarcuatefolds up to 200 m long and 0.5 m in amplitude
that developedwithinlagoonalmuds that were overriddenby
dunes 15-30 m high. McKee et al. (1971) experimentally
producedsmall-scalewarps, folds, andoverthrustsin avalanch-
ing sand, but noted that "no contorted structures caused by
tensionalor compressionalstresses normallyoccur in saltated
deposits.... " There are, furthermore,fundamentaldiffer-
ences in formbetween the productsof lateralcompressionand FIGURE 7-SEM image of sand grains with detritalclay coatings and
the SandHillsdeformationstructures.The circularto ovalplan, bridges. Sample is a "crumb"of recently deposited, but cohesive
lack of directionalasymmetry,and steep marginsof the Sand surface sand collected fromthe side of a vertical-sidedcow track in
Hills structuresgive them a "punched-in" appearance,clearly the central Sand Hills.
indicatingthat they were formed by vertically, rather than
laterallydirectedstress.

The Substrate
Trackmorphologyindicatesthat trackswere made in cohe-
sive sand. Coarse-grainedsedimentsoverlyingtruncatedlam-
inae (Fig. 4a) are lag deposits that filled vertically walled
depressions.Silts preservedin similarpositionsrepresentdust
thatwas trappedandprotectedby concavities.Tracksmadein
moist, well-sortedsand (wetted sand of McKee et al., 1971)
have steep walls, whichcan be maintainedas long as the sand
remainsmoist (Lewis andTitheridge,1978, fig. ld). Were the
Sand Hills tracks formed in moist sand and buried before
drying?Observationsof moder cattle tracksin the SandHills
suggest an alternativehypothesis. Unlike beach sand, dune
sand fromthe SandHillscontainsas muchas 4% silt and clay
(Ahlbrandtand Fryberger, 1980, p. 21). The clay fractionis
smectite, occurringas thin, detrital coatings on sand grains
(Fig. 7; Ahlbrandtand Fryberger, 1982, fig. 21d). Such grain
coatings are deposited by water moving throughthe vadose
zone; they cansurviveconsiderableeoliantransport(Walkeret
al., 1978; Walker,1979).
In the Sand Hills duringAugust, 1985, vertical-sidedcattle
trackswere abundantin wind-rippled,cohesive surface sands
with a moisturecontent less than 1%. "Crumbs"of this cohe-
sive sand collected in the field were found to retain their
structureeven after 24 hours of oven dryingat 40?C. Using
samples of loose sand from the Sand Hills and distilledwater,
cohesive sand "crumbs"identicalto those collected in the field
can be producedin the laboratorywith a single wetting/drying
cycle (Fig. 8).
In the field, cohesive surfacesands are relativelyresistantto
wind erosion. Tracks made within such sands have a much
higherpreservationpotentialthan tracks producedin mobile,
cohesionlesssands that have not been moistened(Figs. 9, 10).
This observationmay explain the prevalence of tracks with
verticalsides in Holocenestrataof the SandHillsand suggests
that clay-coateddune sandis an especiallysuitablemediumfor
the preservationof tracks. In their study of the PermianLyons
Sandstone, Walkerand Harms (1972) hypothesizedthat thin
layers of clay deposited duringcalm periods between sand-
drivingwindsmayhave allowedthe preservationof smalltracks
and raindropimprintsmade in dry sand. Walker(1979) later
showed the importanceof graincoatings composed of detrital
clays to the reddening of eolian dune sands. With further
CENOZOIC
HOOFPRINTS 145
I

FIGURE 8-Crumbs of cohesive sand (left) producedin the laboratory FIGURE 9-Bison tracks in clay-coated eolian sand containing less
when loose surficialsand fromthe Sand Hillswas placed in a paper than 1% water (by weight), FortNiobraraNationalWildlifeRefuge,
cup, saturated with distilled water, and dried in an oven at 40?C. near Valentine.Knifeis 15 cm long.
Cohesionless sample on right was ultrasonicallycleaned and wet
sieved to removeclays priorto identicaltreatment.Originalsample is
a moderatelysorted, nearly symmetricalfine sand (Mz = 2.21; Holoceneage for the deposits; the largest cloven-hoofedmam-
ua=0.662; Skl=.088) containing 1.3% clay, 10 YR7/3 (Munsell mals of the Holocene-bison-are capableof producingall the
ColorChart). tracks so far observed.
Paleoecologicalconsiderationsalso suggest that these tracks
were made by bison. Archaeologicas well as paleontologic
petrographicwork andlaboratoryexperiments,it may be pos- evidencefromseveral GreatPlainsandMiddle
sible to assess the role (if any) of detritalclay coatingsin the sites links bison to semiarid RockyMountain
grassland environments in which
preservationof the delicate surface traces foundin Paleozoic sandy sediments have been subjected to of eolian
eoliansandstones. episodes
transport.Not only have bisonbones been recovered fromthe
Track-Makersand Paleoenvironments Sand Hills (Ahlbrandtand Fryberger, 1980) and from dune
The size and morphologyof the tracks, the probableage of sands at many other localities, but at the 10,000-year-old
in
the Sand Hills, the known mammalianfossil record, and pa- Caspersite east-centralWyoming,paleo-Indiansused para-
leoecologicalarguments strongly suggest that bison are re-
sponsiblefor the tracks. The tracksof modem bison (Fig. 10)
are nearlyidenticalin size andformto those preservedwithin
the SandHills.
+ , E4a g S F F; r .. V.
Untilrecently, students of the Sand Hills had placedmajor
dune formationin either the early or late Wisconsin(Pleisto- b . r. '<*.. , t v f ,.- y
cene). Fluvialdeposits fromthe east-centralSandHills, which l AK~~~~~~~~~~~~~~~~--
AW w
are now knownto underlieas muchas 40 m of dunesand,have,
10 radiocarbon dates from 8410 to
_ - r A
~ v
however, yielded ranging
3000 yrs. B.P., indicatingthat the dune field is primarilyof
ii~G-
Holocene age (Ahlbrandtet al., 1983). This view has been
challengedby Wright et al. (1985), on the basis of radiocarbon ii ' ;' (
X
t h
dates and pollen from five interdune lake deposits in the t P d
d
S? r.
northernandwestern SandHills. At the Calamusdam site, all / 44.'
ijP*pr? S1. $
f P O i
trackslie above a peat depositthathas been radiocarbon-dated 4 e r x / /

at 7260 ? 90 yrs. B.P. (J. Swinehart,unpublisheddata;Beta- .~~~~~~~~~~~~~~~~~~~~~~~.

11621); most tracks are above an organic-richinterdunede-
' ^>
r a'
"i
posit dated at 3450 + 110 yrs. B.P. (ibid, Beta-11622). A ' Pi. .ti" '*
PL I
X rsr.

diverse assemblage of large herbivores, includingcamels, L,

mammoths,horses, and sloths roamed the Great Plains

the Pleistocene;of these, only camels producetracksthat are
during 3'??2,
''?.'t
C 's
C

at all similarin size and shape to those describedhere. The

youngest radiocarbondate for a North Americanfossil site FIGURE 10-Loose sand coveringa deposit of cohesive sand, which
containingcamel bones is 8240 + 960 yrs. B.P. (Mead and contains numerousvertical-sided cattle tracks. CentralSand Hills
Meltzer, 1984). All track observationsare consistent with a near Thedford.Shovel in backgroundfor scale.
146 LOOPE

50 ft-15m -U boliceoliandunes as naturaltraps for bison procurement(Fri-

son, 1974).
The abundanceof tracks in these sediments seems difficult
to reconcile with the lack of root traces below track-bearing
laminae. Were the tracks produced during transient move-
ments of nonresidentherds?The wide spacingof tracks along
beddingplanes and the close vertical spacingof track-bearing
0-0 laminae argue against infrequent mass-migration events.
McKee and Bigarella(1972) observed that small roots pene-
c
tratingdunesandfollowlamination,therebyleavingfew traces.
Trackabundanceand distributionsuggest that semipermanent
-
water-perhaps located in interdune areas-and vegetation
C were availableto resident individualsor herds while the dunes
t
were activelymigrating.
C
OLIGOCENEOF SCOTTSBLUFF NATIONAL
MONUMENT
V
0 Stratigraphicand SedimentologicSetting
c
0O Nonmarinerocks of mid-Tertiaryage are well exposed and
V 0 easily accessible at Scotts BluffNationalMonumentin west-
LU ernmost Nebraska (Fig. 1). These rocks contain abundant
volcaniclasticmaterial,which, together with epiclasticdebris
shed eastwardfrom the Rocky Mountains,blanketedthe sur-
face of the GreatPlains(Stanley,1976; Swinehartet al., 1985).
At Scotts Bluff,siltstones of the BruleFormation(WhiteRiver
Group) are unconformablyoverlain by 27 m of very-fine-
grained,horizontallystratifiedsandstones of the Gering For-
mation(ArikareeGroup)(Fig. 11). Structuresinterpretedhere
vv9 as vertebratetracks are restricted to the lower two-thirdsof
the GeringFormation.The Geringis overlainby about65 m of
9 E large-scalecrossbeddedto massive sandstones of similartex-
0
ture (MonroeCreek-Harrisonunit,Fig. 11). A volcanicash bed
9 T I-
near the base of the Gering at Scotts Bluff has yielded a
/ (/ ) COt
radiometricdate of 25.6 m.y. (Everndenet al., 1964), placing
- o these basal Arikaree rocks within the Oligocene Series
cn: (Harlandet al., 1982). Figure 12 shows the locationof features
0. describedand illustratedin this paper.
a At Scotts Bluff, the Geringis horizontallybedded; the only
channelsobserved duringthis study are a few tens of centi-
vv T meters deep. Small-scale, steeply dipping cross-lamination
:--.V jL _*1 (Fig. 13), andplanarlaminationexhibitingpartinglineation,are
I widespreadthroughoutthis stratigraphicintervaland are clear
evidence of fluvialdeposition(Stanleyand Fagerstrom, 1974).
Earlydiageneticfeaturesin fluvialsedimentscommonlypro-
vide importantclues to depositionalprocesses andpaleoclimate
(Collinson, 1978). Rosettes of calcite-cemented sand com-
posed of discoids up to 8 cm in diameterare present at four
separate stratigraphicintervals within the Gering (Figs. 11,
14). The morphologyof the discoidsandrosettes is identicalto
that of modem gypsum sand crystals described by Cody
.' . I Massive (1979). Afterburial,as the gypsumwas dissolvedby less saline
:.
' sand sand groundwater,replacementof gypsum by calcite was probably
facilitatedby the commonness of the calciumion. The thick
I I Massive :
* volcanicash bed near the middleof the Gering(Fig. 11) andthe
I II siltstone I .|Horizontally-
stratified sand
FIGURE 11-Stratigraphic section and interpretationof depositional
Concretions Silicic ash processes; Scotts Bluff National Monument (from Swinehart and
C-rv Loope, in press). g=evidence of gypsumcrystallization.
 HOOFPRINTS
CENOZOIC 147

21b

0 I
I J
Scale
FIGURE 14-Calcite-cemented sand, pseudomorphousafter gypsum
FIGURE 12-Approximate map patternof contact between BruleFor- "desertroses," middlepartof GeringFormation.Knifehandle is 9 cm
mation (WhiteRiverGroup)and GeringFormation(ArikareeGroup);
from Scottsbluff South Quadrangle,4400-foot contour. Numbers long.
show locations of text figures.
uppermostone meter of the Brule Formationcontainabun-
dant discoids, 0.4 to 8 cm in diameter,that are composed of
calcitespar. Withinthis finer-grainedmaterial,gypsumcrystal
growthpushed the matrixaside to form relativelypure crys-
tals; after dissolutionof gypsum, voids were filledby calcite.
Gypsumcrystals of similarsize and morphologyto those re-
corded in the strata of the study area are today confinedto
inlandand coastal sabkhas, where the groundwatertable ap-
proaches the land surface and undergoes evaporation(Wat-
son, 1983). According to Watson, such materials are re-
3 (Frybergeret al., 1983, fig. 23a; Simpsonand Loope, 1985).
Clr_ZI LY
stricted to environmentswhere annualrainfallis less than 200
mm/yr and where there is a monthly excess of evaporation
over precipitationthroughoutthe year. Evidence of growthof
evaporiteswithinthe relativelypermeablefluvialsands of the
study area strongly suggests that the strata were deposited
by ephemeralstreams.
Several lines of evidence indicate that a portion of the
Arikaree Groupin eastern Wyomingand western Nebraska
accumulatedwithinan eolian dune field (Bart, 1977; Stanley,
1980; Hunt, 1985; Swinehartand Loope, in press). Withinthe
Monroe Creek-Harrisoninterval at Scotts Bluff, starting at
about 10 m above the highest occurrence of tracks, inverse
gradingand wind-rippleforesets (Hunter, 1977) occur in hor-
izontallybedded sands and withinwedge planarcrossbed sets
up to 1.7 m thick. Where the crossbed sets are overlainby
horizontalwind-rippledeposits, the interveningboundingsur-
faces commonlyhave an irregular,"corrugated"appearance
with localrelief up to 10 cm (Fig. 15). Analogoussurfaces are
commonin moder interduneareas where wind erosion has
etched moist, wet, or evaporite-cemented,crossbeddeddune
sands into strong relief (McKee, 1966, pl. VII, c and d;
Fryberger et al. 1983, p. 298). Trenches dug in moder
interdunesreveal crossbeds with wavy or "corrugated"upper
boundingsurfaces, overlainby flat-beddedinterdunedeposits

I
L:
,
Stanley(1980) has noted that physicaland biogenicstructures
?r----cikCi?T P. of the mid-Tertiaryeolianstrataof the GreatPlainsbear many
9_?;rF?T.L=,c-i,
L similaritiesto those withinthe Holocenesands in the Nebraska
.xz
7C?fi
CIPh
Sand Hills. Traces of invertebratesand plantroots are espe-
cJ
6LLCYlk S I ciallyclosely analogous,suggestingto Stanleythat habitatsand
;,? -lj climaticconditionswere very similar.The absence, however,
FIGURE13-Subaqueous climbing-rippledeposits and concave-up of vertebrate tracks in the eolian strata of the Arikaree
deformationstructures, near middle of GeringFormation.Head of Group-both at Scotts Bluff and at the Bear Creek locality
hammeris 18.5 cm long. describedby Bart (1977)-indicates that, unlikethe Holocene
 148 LOOPE

25-

20- N=104

u 15-

O0
u-
5-

2 4

TRACK DIAMETER (CENTIMETERS)

FIGURE 17-Size-frequency data for deformationstructures inter-
FIGURE contact pretedas tracks in GeringFormation,Scotts Bluff. Note that largest
15-Eolian interdunedepositsoverlyingirregular
withcross-stratified
dunesands, MonroeCreek-Harrisoninterval, tracks are largerthan those of Holocene bison (Fig. 5).
summitto museumtrail. Differentialwinderosionof lightlyce-
mentedordampcross-strata tookplaceininterduneareaadjacentto
stoss side of dune.Notrackswereobservedin these eolianstrata.
closely spaced deformationstructures are visible. Some of
these structures have vertically walled infillingsand bilobed
SandHills, the mid-Tertiarydunefields either didnot harbora lower portions (Fig. 19). As in the Sand Hills, tracks were
populationof large vertebrates, or lackedconditonsfavorable producedin relativelyfirmbut compactiblesand. None of the
for the preservationof their tracks. tracks so far observed suggest that the substratewas "quick"
at the time of deformation.Accordingto Simons et al. (1961),
Descriptionand Interpretationof Tracks horizontallylaminatedsands deposited duringupper-flow-re-
gime conditionsare firm relative to sediments deposited by
Concave-updeformationstructures that closely resemble avalanching on the lee side of ripplesor dunes. For most tracks,
the Sand Hills (Holocene) bison tracks are common in the due to uniformityof grain size and to disruptioncaused by
GeringFormation(Figs. 13, 16, 19). Deformationsinthe Gering burrowingand evaporite diagenesis, it is difficultto discern
vary in apparentdiameterfrom4 to 22 cm (Fig. 17). Because whether the parent sediment was originallyparallelor cross-
these rocks are poorlyindurated,bedding-planeexposures are laminated.Therefore, no attempt has been made to compare
small. On the undersidesof overhangingledges, however, the trackdepth in the two types of strata.
circularto oval plan of the deformationstructures is easily In contrastto the Holocenehoofprints,the Oligocenetracks
observed. An extensive search of such exposures revealed appearin sedimentsthatalso show some evidence of physically
several distinct trackwayscomposed of three to five aligned induced deformation:convolute laminationappears in some
tracks (Fig. 18). In vertical outcrops, isolated, paired, and exposures (Fig. 20). In nearlyallcases, however, biogenicand

I, _r- ; M

' '-, ',ts

. '- X

FIGURE16-Large, closely spaced tracks at the tops of numerous, thin sediment packages, near base of Gering Formation.
CENOZOIC
HOOFPRINTS 149
I

FIGURE18-Overhanging ledge revealing four aligned, bilobed FIGURE 19-Cross-sectional view of small, bilobed track made in
tracks, middle partof GeringFormation. cohesive sand by small entelodont or camel; middle part of Gering
Formation.Coin is 1.9 cm in diameter.

physicallyinducedstructurescan be confidentlydifferentiated (LaporteandBehrensmeyer,1980, fig. 4a). Anotheris that the

on the basis of scale, continuity,or three-dimensionalgeome-
try.
A diverse assemblageof large mammalsis knownfromlate
Oligocenerocks of the Great Plains. The distinct polymodal
natureof the size-frequencydistribution(Fig. 17) suggests that
the tracksof severalspecies are preservedin the Geringsands.
The moder ungulateorders, PerissodactylaandArtiodactyla,
first appeared in the Eocene (Romer, 1966). By the late
Oligocene, diversificationof artiodactylsubgroups was well
under way. Representative taxa of six artiodactyland four
perissodactylfamiliesare knownin Geringsedimentsof Wildcat
Ridge, southeast of Scotts Bluff (Swisher, 1982). A single
fortuitousbedding-planeexposure (Fig. 21a) indicatesthat at
least some of the largest trackswere producedby a two-toed
animal. Morphologicallysimilar tracks from Oligocene and
Miocene rocks have been described by Robertson and
Sternberg (1942), Chaffee (1943), Bjork (1976), and Dema-
thieu et al. (1984). Of the two-toed members of the Gering
faunal assemblage, only entelodonts were large enough to
makethese tracks(R. M. Hunt,personalcommunication). The
small,bilobedtracks(Figs. 18, 19) were probablyproducedby
smallerartiodactylssuch as camels.
It is difficultto dividea horizontallybedded sequence com-
posed of very-well-sortedsediment into distinct depositional
packages. Traces of invertebrates provide clues in marine
populationdensity of large vertebrates was very low. A third
possibilityis that evaporiticsurface crusts formed after each
pulse of sedimentation.Early evaporite diagenesis may have
quicklymade the sandy substrate less compactibleand thus
unsuitablefor preservationof tracks. If this was the case, only
a very smallpercentageof the vertebrateactivitythatoccurred
here maybe recordedby the tracks.The preservationpotential
of tracks or other surficialtraces in sandy fluvialsequences
would seem to be quite low because of their vulnerabilityto
scour. Earlyevaporitecementationmay partiallyexplainboth
the preservationof tracksand, conversely, the rarityandsmall
scale of channelsin these ancientfluvialstrata.
CONCLUSIONS
1. HoloceneeolianandOligocenefluvialsedimentsof Nebraska
contain abundanttracks of large vertebrates. The most
c i
n_.r -?-?pr b ;r;?
-s-.-- - ka .
71- - CJii;'.?i-regFI. j-a-l;?
inh

sequences (Howard,1978); tracksof terrestrialvertebratesin

the Geringcan be utilizedin a similarway. The trackdistribu-
tion at Scotts Bluff mirrors that of the Sand Hills: laterally
scattered tracks are typicallyseen at closely spaced vertical
intervals.If tracks are assumed to markboundariesbetween
depositionalevents, the distributionof tracks indicates that
streamfloodsdepositedpackagesof stratabetween a few cen-
timeters to over three meters in thickness. Thoroughlytram-
pled horizons(Fig. 16), as mightbe expected alongdiastems,
are relativelyrare. Why are the tracks that markthe tops of
sediment packages so widely spaced? One possibilityis that FIGURE 20-Convolute lamination(nonbiogenic),middlepartof Ger-
time intervals between depositionalevents were very brief ing Formation.Note consistent asymmetryof folds.
150
4I-? ..
'Wu
, have enhancedthe preservationpotentialof individualtracks
4
, .
.
.:" i.-'O I
ap . 41
All-~C4
"YW'T
r 40L
FIGURE21-Tracks of large entelodonts. A) Small, overhanging ledge
revealing two large tracks (12 and 16 cm in length). Both tracks are
cloven in front; large track was made by front foot. B) Cross section
of asymmetric pair of tracks. Deeper track on right probably made by
front foot. Middle part of Gering Formation.
obvious manifestationof the tracks is the downwarpingof
laminaeseen in verticalsection.
2. Although tracks may superficiallyresemble nonbiogenic
soft-sedimentdeformationstructures, their isolationalong
bedding planes, circularto oval plan, and restricted size
distributionaid in their recognition.Convolutebeddingof
physical origin is generally restricted to initiallyporous,
rapidlydepositedsediments.
3. TrackswithinHolocenestrataof the SandHillswere made
in clay-coatedsands deposited by migratingwind ripples.
Resident bison that were sustainedby food and water re-
sources made the tracks while the dune field was active.
4. Oligocenerocks at Scotts Bluffare preservedat the tops of
thin packages of fluvial deposits. Abundantevidence of
evaporite precipitationsuggests deposition by ephemeral
streams. Size-frequencyplots indicate that tracks were
made by several differenttypes of ungulates;entelodonts
producedthe largesttracks.Cementationby evaporitesmay
LOOPE
by preventingscour and intense trampling.
ACKNOWLEDGMENTS
J. B. Swinehartprovidedtimely field trips and lively discus-
sions. R. M. Hunt and M. R. Voorhies freely shared their
knowledgeof Cenozoicfaunasandenvironments.I thankthese
three Universityof Nebraskacolleagues for their enthusiasm
and editorialassistance. DonaldBairdof PrincetonUniversity
reviewedthe manuscriptandprovidedadditionalreferences on
mammaltracks. Larry Cast of the Bureau of Reclamation
helped with access to the Calamusdam site. Mike Leite's
darkroomskills improvedmany of the photographs;Kit Lee
helpedwith the SEM. Acknowledgmentis made to the Donors
of the PetroleumResearch Fund, administeredby the Ameri-
can ChemicalSociety, for supportof this research.
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