Paleontological Society
Blancan Lagomorphs and Rodents of the Deer Park Assemblages, Meade County, Kansas
Author(s): Robert A. Martin, James G. Honey, Pablo Peláez-Campomanes, H. Thomas Goodwin,
Jon A. Baskin and Richard J. Zakrzewski
Source: Journal of Paleontology, Vol. 76, No. 6 (Nov., 2002), pp. 1072-1090
Published by: Paleontological Society
Stable URL: http://www.jstor.org/stable/1307124 .
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J. Paleont., 76(6), 2002, pp. 1072-1090
Copyright ? 2002, The Paleontological Society
0022-3360/02/0076-1072$03.00
BLANCAN LAGOMORPHSAND RODENTSOF THE DEER PARK
ASSEMBLAGES,MEADE COUNTY,KANSAS
ROBERT A. MARTIN,' JAMES G. HONEY,2 PABLO PELAEZ-CAMPOMANES,3 H. THOMAS GOODWIN,4
JON A. BASKIN,5 ANDRICHARD J. ZAKRZEWSKI6
'Departmentof Biological Sciences,MurrayStateUniversity,Murray,Kentucky42071, <[email protected]>,
2GeologySection,Universityof Colorado,Boulder80309-0315,
of Paleobiology,NationalMuseumof NaturalHistory,C.S.I.C.,Jose GutierrezAbascal2, Madrid28006, Spain,
3Department
of Biology, AndrewsUniversity,BerrienSprings,Michigan49104,
4Department
of Biology, TexasA & M University,Kingsville78363, and
5Department
of Geosciencesand SternbergMuseumof NaturalHistory,FortHays StateUniversity,Hays, Kansas67601
6Department
ABSTRACT-A new collectionof lagomorphsandrodentsfromthe Deer ParkB local fauna(1.f.)of MeadeCounty,Kansasis described
and comparedwith othersmall mammalassemblagesof the MeadeBasin, includingthe underlyingDeer ParkA 1.f.Deer ParkA was
correctlyassignedby Hibbardto the Blancan,bridgingthe gapbetweenearlierBlancanfaunassuchas Fox Canyonandthe late Blancan
Sanders1.f.Recentfieldworkindicatesthatthe DeerParkquarriesmaylie in the RexroadFormation,ratherthanin the BallardFormation
as previouslyassumed.The geology and extinctmammaliancontingentat Deer Parksuggest that the lower horizonof Deer ParkA
was an active springthat graduallyturnedinto a marshyenvironmentduringDeer ParkB time. The rodentsof Deer ParkB are
indicativeof an open prairieecosystemthatmighthave been somewhatmore aridthanthatof southwesternKansastoday.
IN1997 R. Martin, J. Honey, and P Peliez-Campomanes initi-
ated a long-term field and analytical project to study rodent
community evolution during the late Cenozoic in southwestern
Kansas. In addition to prospecting for new localities to fill in gaps
in the Pliocene and Pleistocene sequence previously established
by the late C. W. Hibbard and his students (e.g., Hibbard, 1972;
Woodburne, 1961; Zakrzewski, 1975) we resampled many of Hibbard's original localities (Martin et al., 2000). This paper represents the first in a series of faunal studies from this ongoing field
program.
The Deer Park locality was the first site that Hibbard worked
in the Meade Basin. In 1936, Dr. M. K. Elias, then of the Kansas
State Geological Survey, brought some fossil mammal teeth to
Hibbard at the Kansas University Museum (now the Museum of
Natural History). Workers at a Civilian Conservation Corp (CCC)
camp on Meade County State Park property, later to be Hibbard's
field headquarters for many years, had found fossil teeth in the
park and were distributing them as souvenirs. Hibbard and a field
party from the University of Kansas collected at the locality, subsequently known as Rexroad Loc. No. 1, in the summer of 1936,
but at the time they thought the deposit had been gutted, and only
collected from the surface. They again surface collected there in
1937 and 1944, and in 1954 they dug two trenches, about 19 m
apart, into the base of the east side of the small ravine near where
the CCC workers had been digging. Hibbard's two spoil piles
currently identify the sites. The locality is found in Meade State
Park, at the west edge of the SE1/4sec. 15, T33S, R29W. According to Hibbard (1956, p. 170), the fossils ".... are found in floursand pockets and tubes developed in the silty clay by an old, nowdry, artesian-spring system." In addition to recovering large vertebrates, they dry-sieved the fine sand over window screen for
smaller fossils.
Although there is no accompanying explanation, Hibbard
(1938) included description of fossil remains from another locality, Locality 2 (=Rexroad Loc. 2), along with his initial description of fossils from Deer Park (=Rexroad Loc. 1). At that time
Hibbard considered the fossils collected from Loc. 1 to be part
of his "Rexroad fauna," and only later (Hibbard, 1949) did he
formally separate them as the distinct Deer Park local fauna, although he recognized earlier that they might be younger than
material from the other Rexroad localites (Frye and Hibbard,
1941). Loc. 2 is on private land less than 0.5 km from Loc. 1.
To make matters more difficult, there are two localities in the
pasture where Loc. 2 is found; one is Loc. 2, the other Loc. 2a
(Woodburne, 1961), about 50 m to the southwest of Loc. 2. Hibbard (1956) later described the fossils from Loc. 1 separately, but
there has been considerable confusion regarding those that were
recovered from Locs. 2 and 2a, compounded by the fact that the
holotype of Pliopotamys meadensis (=Neondatra kansasensis),
listed in Hibbard (1938) as from Loc. 2, was apparentlyrecovered
from Loc. 1 (Hibbard, 1956; p. 175). Both Rexroad Loc. 2 and
2a appear to be stratigraphically higher than the Deer Park sites.
The fossil fauna from Rexroad Loc. 2 was taken by Hibbardfrom
a narrow flour sand tube mined to a depth of more than three
meters. The excavation site of Loc. 2 is no longer visible on the
upland prairie surface, and no further attempt was made to find it.
In the following sections we describe our current view of the
stratigraphic position of the Deer Park assemblages, laboratory
analytical methods, and the species accounts for the lagomorphs
and rodents. In Table 1, we also present some new records from
the University of Michigan, Museum of Paleontology collections
for the Bender lb and ic l.f.s and a few records from Rexroad
Loc. 2a, based exclusively on a small collection made during the
summer of 2001. We conclude with a discussion relating the Deer
Park l.f.s to other Blancan l.f.s from southwestern Kansas and
also consider the ecological setting of the Deer Park quarry area
during the time of deposition.
GEOLOGIC SETTING
As noted above, Hibbard's excavations were into flour-sand
tubes invading reddish sediments at the base of a ravine in Meade
County State Park (Hibbard, 1956). Our excavations were made
into a horizontal gray-green sandy clay unit directly overlying the
red sediments reported by Hibbard (see below). Hibbard'soriginal
locality is thereby recognized as Deer Park A, and ours as Deer
Park B. Fossils recovered from these sites are designated the Deer
Park A and B 1.f.s, respectively. In the summer of 2000 we dug
a stairstep trench from the base of our excavation down into the
base of the ravine. We determined that the upper gray-green unit
grades into the lower one with no discontinuity. We did not encounter flour-sand tubes; Hibbard may have mined them out, or
they may exist lateral to our test trench. Our first excavation in
1997 encompassed an area approximately 3 m wide by 1 m deep.
We took 100 sacks from this pit, representing about two tons of
matrix. This sediment was wet screened in the field with 0.5 mm
1072
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MARTINET AL.-BLANCAN RODENTS AND LAGOMORPHSFROM KANSAS
1073
1-Measurements (in mm) of Geomys quinni, G. jacobi and G. minor lower fourthpremolarsand upperincisors from Kansas and Nebraska.Numbers,
from top to bottom in each cell, are: numberof specimens, mean, observed range and standarddeviation. See text for measurementdescription.
TABLE
WA
G. quinni
Deer Park B
G. quinni
Sand Draw
G. jacobi
Rexroad Loc. 3
G. jacobi
Wendell Fox
Pasture
G. minor
Deer Park B
G. minor
Rexroad Loc. 3
10
1.83
(1.61-1.98)
0.11
12
1.92
(1.72-2.18)
0.14
18
1.91
(1.75-2.11)
0.11
9
1.70
(1.57-1.81)
0.09
2
1.17
(1.16-1.17)
15
1.20
(1.02-1.47)
0.12
C1l
10
0.11
(0.06-0.22)
0.05
12
0.14
(0.07-0.41)
0.09
19
0.11
(0.03-0.19)
0.04
10
0.11
(0.07-0.14)
0.02
4
0.09
(0.06-0.17)
0.05
15
0.06
(0.02-0.09)
0.02
sieves. In the summerof 1999, we washedan additional50 bags
of matrixfrom the same quarry.
Hibbardinitially thoughtthat the sedimentsfrom which the
Deer ParkA 1.f. was collected belongedto the RexroadFormation, but later (Hibbard,1949), he includedthem in the Missler
Memberof the Meade Formation.The term BallardFormation
was subsequentlyselected as a replacementname for the Meade
Formation(Hibbard,1958). The type locality of the BallardFormation is on the formerBig SpringsRanch, in Meade County,
less than 10 km northof Deer Park.In October,1999 we measuredthe stratigraphic
positionof the Deer Parkquarriesrelative
to the surroundingsediments,and our resultsare providedbelow
in a measuredsection. Two gravels crop out in the Deer Park
area;a lower unit that is a conglomerateat its base and unconsolidatedhigherup, andan upper,unconsolidatedgravel.Because
of the unconsolidatednatureof these gravelsandthe grasscover,
theirpositionscan only be approximated.
The unconsolidatedupper part of the lower gravel is encounteredbelow the quarries
about 15 m to the south.About 1.0 m of the lower conglomerate
is exposed approximately20 m west of the Deer Parkquarries,
at the foot of the ravinein which the quarriesare located.There
are about 15-16 m of sedimentsbetween the top of the lower
gravel and the contact with the upper gravel in the Deer Park
area.The sedimentsbetweenthe two gravelsare mostly covered
by grass, and we did not attemptto characterizethe few areas
where they are exposed. The Deer Parkquarrieslie about 2 m
above the estimatedtop of the lower gravel.We assumethatthe
uppergravelis the StumpArroyoMemberof the CrookedCreek
Formation,but we are currentlyuncertainof the identityof the
lower gravel. It has in the past been equatedwith the Angell
("MeadeGravels")Memberof the Ballard("Meade")Formation
(Hibbard,1956; Stevens, 1965; Martinet al., 2000) which, accordingto Hibbard(1958), overliesthe RexroadFormation.However, in the summerof 2001 we determinedthat RexroadLocs.
2 and 2a and apparentlyalso the classic RexroadLoc. 3 (type
RexroadFormation)lies above a similarlooking gravel,mapped
as withinthe BallardFormation(Woodburne,1961,or specifically
as the basal (Angell) gravel of the BallardFormation(Stevens,
1965). Both the RexroadLoc. 3 and Deer Parkassemblagesare
thereforeyoungerthan local faunasfrom the Bender lb and Ic
Wp4
12
2.35
(2.14-2.49)
0.06
12
2.57
(2.29-2.85)
0.17
19
2.54
(2.21-2.98)
0.18
10
2.29
(2.09-2.37)
0.08
4
1.46
(1.41-1.51)
0.04
15
1.54
(1.23-1.78)
0.14
Lp4
11
3.12
(2.71-3.78)
0.28
12
3.55
(3.26-3.94)
0.22
18
3.35
(3.00-4.04)
0.30
10
3.09
(2.82-3.42)
0.17
2
1.69
(1.49-1.88)
0.28
15
2.13
(1.81-2.42)
0.19
WI
33
3.18
(2.71-3.49)
0.24
7
2.97
(2.78-3.20)
0.17
10
3.18
(2.65-3.43)
0.25
10
2.97
(2.61-3.33)
0.22
12
1.84
(1.50-2.07)
0.17
10
1.95
(1.72-2.35)
0.22
and Hibbard'sUM-K1-59 (=our Hornet)localities,as the latter
localities lie stratigraphically
below this lower gravel. Thus, the
position of RexroadLoc. 3 above this lower gravel, which has
previouslybeen identifiedas the Angell Member,throwsserious
doubton the correlationof this gravelwith the type Angell Member on the Big SpringsRanch,andpossiblyon the validityof the
BallardFormationas a rock unit distinctfrom and overlyingthe
RexroadFormation.Because the type area of the BallardFormationon the formerBig SpringsRanchis currentlyunavailable
to scientists,resolutionof this particularproblemmay take some
time. Furtherdeterminationof the age of the Deer Parkassemblages dependsto a greatextenton the rodentcontingentandwill
be consideredin the discussionfollowingthe species accounts.It
seems likely, based on these preliminaryconsiderations,thatthe
Deer Parklocality residesin the RexroadFormation,as Hibbard
originallysuggested.
Rodentsfromthe Benderlocal faunawere recoveredfromtwo
localities, labeled lb and Ic by Taylor (1960). Locality lb is
underliesthe lowergravel,as notaccessible,andstratigraphically
ed above. Locality Ic could not be found duringthe summerof
2001, and is apparentlynow buriedundertons of slumpedearth.
Accordingto Taylor's(1960) directions,Loc. Ic should be approximatelyat the same stratigraphiclevel as lb, and our field
observationsdid not refutethatpossibility.Species from the two
localities are presentedin Table 1. Referralin the text to the
"Bender1.f."impliesa tentativecombinationof thesefaunallists.
As will be discussedbelow underPaleoecology,some physical
modificationof fossils recoveredfrom the Deer ParkB quarry
probablyresultedfrom mineralpercolationand biologicalinfluences (e.g., plant root formation)in the modernsoil, which is
developedthroughpartof our quarry.Althoughthereare numerous soil classifications,most ecologists recognizefive basic horizons, labeledby Ricklefs and Miller (1999) as O, A, E, B, and
C. Most of the organicmaterialis locatedin the O and A levels.
In E, B, andC, leachingandmineraldepositionis seen. Sediments
throughouta profile,if the soil is extensive,arecharacteristically
columnar,or blocky, particularlyif root growthis common.Levels 4-8 of our measuredsection below are presumedto be part
of the modernHigh Plainsshortgrassprairiesoil profile.
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1074
JOURNALOF PALEONTOLOGY,
V. 76, NO. 6, 2002
MEASURED SECTION AT DEER PARK QUARRIES
Blancan= 1.8-3.0 Ma, middleBlancan= 3.0-4.0 Ma, andearly
Blancan = 4.0-5.0 Ma.
Institutionalabbreviations
areas follows:MSU = MurrayState
University;FHSM= FortHays SternbergMuseum;UCR = Uni?StumpArroyoMemberof the CrookedCreekFormation
versityof California,Riverside;UMMP = Universityof Michi10. Gravel.............................
not measured gan, Museum of Paleontology;IGCU = Instituteof Geology,
?RexroadFormation
UniversidadNacionalAutonomade Mexico;F:AM = FrickCol9. Coveredinterval........................
12.2-13.1 lection,AmericanMuseumof NaturalHistory.All specimensre8. Organiclayers(O andA horizons)of modernsoil; gray
portedfrom Deer ParkB bearFHSMVP numbers.
to darkbrown.............................
0.30
7. Sandy,silty mudstone,buff; blocky with vertical
SYSTEMATIC PALEONTOLOGY
fractures.................................
0.33
we
Although recognizethatall speciesbasedon fossil materials
6. Clay, brownish-gray,sandy (fine grained),containing
thin CaCO3laminae;blocky with verticalfractures 0.30 are morphospecies,we have tried in our Meade Basin work to
represent,as faithfullyas possible,the biologicalrealityof species
5. Sandyclay and clayey, very fine-grainedsand;grayishcontinuitythroughtime. In this treatmenta speciesis synonymous
with
scattered
green
light gray mottling (probably
with a lineage, and no matterhow much change occurs in that
some
limonitic
staincontains
orange-brown?
CaCO3);
lineage
only one species is recognized.The rationaleandphilos0.15
ing. Sourceof Deer ParkB l.f .................
08 ophy of this approachhavebeen discussedin detail(Martin,1993,
4. Calichezone, discontinuous...................
1995; Carroll,1997) and will not be repeatedhere. The reader
3. Sandstone,pinkish-brown,very fine-grained,very claywill
find in the text the use of Martin's(1993) chronomorph,a
massive.
Lower
contains
scattered
caliche
nodules;
ey;
modified
versionof the "lineagesegment"conceptof Krishtalka
partof unit covered;we reporta partialthickness.Apand
(1985). As is a lineage segment,a chronomorphcan
Stucky
A
Park
0.46
source
of
Deer
parent
l.f................
1.52 be an intermediatepopulationin a phyleticsequence,with a par2. Coveredinterval............................
ticularfossil sample informallydesignatedas a referent.It is a
Unknownunit, previouslyidentifiedas Angell Member
somewhatarbitrarydesignation,providedprimarilyfor biostrati1. Conglomerateandsandstone,yellowish,unconsolidated
graphic
purposes.Unlike lineage segments, chronomorphscan
not
measured
above base .................
........
15.3-16.2 overlapin time, to the extent a mosaic patternof evolutioncharTotalmeasuredsection ...............
chronacterizesthe history of a given taxon. Contemporaneous
omorphsare operationallydefined as geographicallyseparated,
MEASUREMENTS AND ABBREVIATIONS
morphologicallydistinctfossil samples within one species that,
when alive, would have presumablyrepresentedpopulationsat a
Dental measurementson the leporids,geomyids,heteromyids, hierarchical
level below subspecies(e.g., the five versus six triarvicolids,and some cricetidswere made with an ocularmicroml populations of modem Microtus pennsylvanicus; Davis,
angle
meter and binocularmicroscope,calibratedwith a 2.0 mm AO
The distinctionbetween extinct subspeciesand contemslide. Most of the remainingcricetidteeth were measuredwith a 1987).
is admittedlysomewhatfuzzy, but in
poraneous
Nikonmonocularmeasuringmicroscopewith digitalreadout.The the case ofchronomorphsthe
identifiedpopulationsalso have a
chronomorphs
Microa
Gaertner
were
measured
woodratteeth
Measuring
using
modem mammalogistswould not
Thus,
chronological
history.
scope. For most sciurid specimens, measurementswere taken
populationsof Microtuspennsylvanicuswith a domifrom calibratedimages capturedthrougha Wild MZ-8 stereomi- recognize
nanceof five triangleson ml as a subspeciesdistinctfromthose
croscopeusing NIH Image.Illustrationsof otherspecimenswere with a dominanceof six triangles,but we can see this transition
made with a cameralucida attachmentto an Olympusor Wild
place in the fossil record,with some overlaprecordedat
M8 binocularmicroscope.Lengthsand widths are greatestmea- taking
various points, includingmoderntime. Chronomorphsare recsurementstaken on the occlusal surface,unless otherwiseindiby a trinomialprecededby a "slash" mark,such as in
cated. Upper case indicates upper dentition,lower case, lower ognized
Ondatra zibethicus Imeadensis.
dentition.Left andright are indicatedby uppercase L and R; T
= triangleon arvicolidmolars.Dentalterminologyfollows White
Order LAGOMORPHA
Brandt, 1855
(1987, 1991) for leporids,Goodwin(1995) for sciurids,Tomida
LEPORIDAE
Gray, 1821
Family
(1987) for the major and minor cusps and Martinand Prince
LEPORINAETrouessart, 1880
Subfamily
(1989) for re-entrantfolds of cricetids,van der Meulen (1973)
ALILEPUS sp.
and Martin(1987) for arvicolids,and Barnosky(1986) for hetFigure2.1
eromyids.Re-entrantangles (synclines)in arvicolidmolarsthat
Material
as
those
to
here
referred
curve anteriorlyare
examined.--FHSMVP-14064,L p3; 14065, R p3.
perpenprovergent;
dicularto the molarlong axis arehorizontal.Reentrantsthatcurve
Discussion.--Two p3s are referred to Alilepus. This genus is
posteriorly,as in the upperdentition,arepostvergent(termspartly recognizedon the basis of the retentionof a relativelyshallow
from Rabeder,1981).
posterior external reentrant(PER) with minimal crenulations
Measurementsof p4s and Is for pocketgophersare as follows: (folding) on its anteriorsurface,in combinationwith a well-deWA = greatestwidthacrossthe anterolophidof p4; Cl = closure, veloped PIR (posteriorinternalreentrant),occasionallyisolated
the widthof the dentineisthmusconnectingthe anterolophidand into an enamel lake. Both specimensare heavily worn, but the
posterolophid;Wp4 = greatestwidth across the posterolophid; AIR (anteriorinternalreentrant)on VP-14064 may have been
Lp4 = greatestlengthof the p4; WI = greatestwidthof the upper isolated in a lake, suggestingaffinitywith A. vagus ratherthan
with A. hibbardi.The latterhas a shallow PIR and has thusfar
incisor.
andHemphillianlocalities
At least informally,the Blancanis often brokeninto "early" been recordedonly fromClarendonian
and "late" intervals(e.g., Lundeliuset al., 1987). These terms (White,1991).The Deer ParkB p3s areconsiderablysmallerthan
have little biologicalor stratigraphic
meaningin the MeadeBasin those of A. wilsoni, namedby White (1991) from the Borchers
sequence,and for purelypracticalreasonswe preferto breakthe 1.f.Lengthsand widthsof VP-14064and 14065 are,respectively:
Blancaninto threeintervalsof roughlyequalduration.Thus,late L = 3.28, 2.93 mm; W = 3.17, 2.76 mm.
Top of Section
Thickness (m)
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MARTINET AL.-BLANCANRODENTSAND LAGOMORPHS
FROMKANSAS
LOCAL
FAUNAS
West, Mexico
Ma
MPTS
LMA
1075
CentralU. S.
RLB 0
Cudahy
SLCB
0
1.0
z
AriesB
Aries A
-----I
Nash 72
2.0
ECasco
1
3
2
Rick Forester
AriesNE
ShortHaul
Borchers
Curtis Ranch
1
HR Margaret
Sanders,SandDraw
Tusker
Deer Park A, B
RexroadLoc.3
Bender lb, ic
Benson
Rexroad
2a
3.0
Hornet
4.0
H
.0
<
z
S
5.0
Wiens, Vasquez
Keefe Canyon,RaptoriC
Ripley
Fox Canyon
Fallen Angel
Saw RockCanyon
Pipe Creek
Buis Ranch,
Devil's Nest, Santee
Mailbox
Hagerman
11
10
12
13
Concha
Verde
Yep6mera
Panaca
14
15
16
17
2-Lagomorphs from Deer ParkB and geomyidsfromthe Deer
FIGURE
Parkassemblagesanda varietyof late Blancanand modernlocalities.
Alilepus sp.: 1) FHSM VP-14064, L p3. Nekrolagusprogressus:2)
FHSM VP-14062, R p3. Notolaguslepusculus:3) FHSM VP-14066,
L p3. Geomys(Geomys)cf. quinni(DeerParkB): 4) FHSMVP-14425,
FIGURE1-Stratigraphic relationshipsand chronologicalsummaryof
R p4; 5) FHSMVP-14428,L p4; 6) FHSMVP-14424L p4; 7) FHSM
Meade Basin fossil localities, includingselect sites from other geoVP-14427 L p4; 8) FHSMVP-14426 L p4. G. quinni(Deer ParkA):
graphicregionsin the UnitedStatesandMexico. LMA = NorthAmer9) UMMP31956(1), Lp 4; 10) UMMP31956(2),R p4. Geomys(Geoican LandMammalAge, MPTS= MagneticPolarityTimeScale (from
mys)
jacobi (RexroadLoc. 3): 11) frombulkp4s UMMP42278, R p4.
=
=
et
Ma
Million
LCB
Lava Creek
Berggren al., 1995),
years ago,
Geomys
(Geomys)bursarius (modern;Meade Co. State Park): 12)
Basin, CTB = CerroToledo Basin; HR = HuckleberryRidge ash.
UMMZ 109932,R p4. Geomys(Nerterogeomys)
cf. minor(Deer Park
MeadeBasin faunalsequencemodifiedfromMartinet al. (2000).
B): 13) FHSM VP-14430,L p4 and 14) FHSM VP-14439 (juvenile),
R p4; G. minor(RexroadLoc. 3): 15) and 16) frombulk p4s UMMP
53924, R p4s. Scale bars (upperfor lagomorphs,lower for geomyids)
= 1.0 mm.
NEKROLAGUSPROGRESSUSHibbard, 1939
Figure 2.2
Material examined.-FHSM VP-14062, R p3.
Discussion.-Nekrolagus progressus is a moderately large lep- and, in the Meade Basin, from the Keefe Canyon and Rexroad
orine with well-developed AER, PER, and PIR on p3. The latter Loc. 3 sites (White, 1991). The Deer Park specimen was broken
forms a large isolated lake, almost directly in contact with the and accurate measurements could not be taken.
internal border of the PER. Additionally, a small but distinct anOrder RODENTIA
Bowdich, 1821
terior reentrant (AR) is present as a groove that is clearly disFamily SCIURIDAEGray, 1821
cernible along the entire anterior p3 face. The Deer Park B p3 is
Figures 3, 4
heavily worn, but the well-incised AR, filled with cement, is obvious in occlusal view (Fig. 2B), and although the lake is not
Numerous species of ground-dwelling squirrels have been depreserved on the occlusal surface, it can be seen from the under- scribed from the Blancan of the Great Plains. Most species are
side of the tooth. Length of VP-14062 = 2.92 mm, W = 2.66 generalized ground squirrels broadly similar in dental pattern
mm.
and identification can be difficult, especially based on isolated
teeth. Metric comparisons on ml-m2 show two size clusters
NOTOLAGUS
Hibbard, 1939
LEPUSCULUS
(Fig. 3). Small species include Spermophilus howelli from the
Figure 2.3
Blancan Rexroad Loc. 3 (type locality) and Fox Canyon (HibMaterial examined.--FHSM VP-14066, L p3.
bard, 1941b; Zakrzewski, 1969); S. meltoni from the late Blancan
Discussion.-Notolagus lepusculus is a relatively small leporid Sand Draw 1.f. of Nebraska (Skinner and Hibbard, 1972); and S.
with a deep, highly crenulated PIR on p3. There are also usually meadensis from the latest Blancan Borchers
1.f. (Hibbard, 1941a).
two crenulated external reentrants. Unlike in N. velox, the AER The Fox Canyon squirrel referred to S. howelli averages noticedoes not connect with the internal reentrantin N. lepusculus. No- ably larger than S. howelli from the type locality (Fig. 3; statistical
tolagus lepusculus is a characteristic Blancan leporine, known tests not performed because samples lump ml and m2) and seems
from Texas (Beck Ranch), New Mexico (Truthor Consequences), best identified as S. cf. howelli.
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1076
JOURNALOF PALEONTOLOGY,
V. 76, NO. 6, 2002
4.0111
1
SDeer ParkSpermophilus
*
E S. boothi
O S.johnsoni
3
/
0 S. mcgheei
•E
E 3.5
-
Ct
$
S. finalyensis
S. dotti
A
S. meltoni,
(n 4) /
S. meadensis
2.5
(n
(n=13)
?i
0-.07
y-0.99x
+ 0.02
E 3.0
" 2.5
"
y=1.28x
S. exroadensis
(RexroadLoc.3;
n=14)
n=14)
S.•/ cf. S. rexroadensis
(FoxCanyon;n=23)
-
S. of. S. howelli(FoxCanyon;n=47)
2.0
2.0
SShowelli(RexroadLoc.3: n=7)
2.5
3.0
3.5
4.0
1mm
Talonid width ml or m2 (mm)
FIGURE
3-Scatterplot of trigonidand talonidwidths of ml-m2s com- FIGURE
4-Representative teeth of Spermophilusfrom Deer ParkB. 1paringlatest Hemphillianand BlancanSpermophilusfrom the Great
2, FHSM VP-14225-R ml or m2 of Spermophilussp. (small) in ocPlainsand fossils reportedfromDeer ParkB. Separateregressionsare
clusal (1) andbuccal(2) views. 3-4, FHSMVP-14242-RM1 of S. cf.
plottedfor clustersof small (S. meadensis,S. howelli, S. cf. howelli,
S. rexroadensisin buccal (3) and occlusal(4) views.
S. meltoni)and large (S. rexroadensis,S. cf. rexroadensis,S. boothi,
S. johnsoni, S. mcgheei,S. finlayensis)Spermophilus.S. dotti was not
includedin regressionsbecauseit is intermediate
betweensize clusters.
A cluster of larger squirrels includes S. rexroadensis from
Rexroad Loc. 3 (type locality) and Fox Canyon (Hibbard, 1941b;
Hazard, 1961); S. finlayensis and S. mcgheei from the late Blancan
Hudspeth 1.f., Texas (Strain, 1966); and S. boothi and S. johnsoni
from the late Blancan Sand Draw 1.f., Nebraska (Skinner and
Hibbard, 1972). Hazard (1961) assigned the large ground squirrel
from Fox Canyon to S. rexroadensis without hesitation, although
this sample appears to average substantially smaller than that from
the type locality (Fig. 3; statistical tests not performed because
samples include both mls and m2s) and is best identified as S.
cf. rexroadensis. S. dotti, known from the latest Hemphillian Buis
Ranch l.f., Oklahoma, appears intermediate between the clusters
(Fig. 3).
The two size clusters differ in the relationship between trigonid
and talonid widths of combined ml-m2s (Fig. 3). Treated as a
group, small Spermophilus show an isometric relationship between these dimensions, with trigonid and talonid widths essentially equal across the observed size range. In contrast, large Spermophilus, as a group, exhibit a significantly narrower trigonid at
smaller sizes (the slopes of regressions for small and large samples were significantly different based on analysis of covarianceF = 5.14, P < 0.05). The intermediate S. dotti follows the isometric relationship exhibited by small species.
At least two species of Spermophilus are represented in the
Deer Park B 1.f., representing both size clusters. One specimen
represents a small squirrel with greatest metric similarity to S.
meadensis and S. howelli from Rexroad Locality 3, whereas seven
specimens, including two measurable ml-m2s (Fig. 3), show
greatest metric similarities to the Fox Canyon sample of S. cf.
rexroadensis or to S. finlayensis (Fig. 3).
Hibbard (1956) described a prairie dog from Deer Park A, Cynomys meadensis, but subsequent work has suggested that this
species was an intrusive C. ludovicianus, the black-tailed prairie
dog characteristic of the region today and present in the Rancholabrean (Goodwin, 1995). Prairie dogs (Cynomys) and prairie
dog-like ground squirrels (e.g., ?Spermophilus cragini from
Borchers) are known from the late Blancan (Goodwin, 1995;
Goodwin and Hayes, 1994), but none of the specimens from Deer
Park B exhibit the derived dental morphology characteristic of
these forms.
SPERMOPHILUS
sp. (small)
Figure 4.1, 4.2
Material examined.-FHSM VP-14225, Rml or m2.
Discussion.-The single, lightly worn lower molar represents
a small squirrel (Fig. 3; L = 1.92 mm, TrW = 2.36 mm, TlW =
2.27 mm). This specimen bears a prominent trigonid pit bounded
by a distinct protolophid and complete metalophid (Fig. 4.1), and
probably is an ml (ml characteristically exhibits a more developed metalophid than does m2 in Spermophilus). A line connecting the lingual surfaces of entoconid and metaconid forms an
angle of 61 degrees with the protoconid-metaconid axis, resembling S. meadensis (64 degrees, n = 8). This is more acute than
typical of S. howelli (68 degrees, n = 12) or a single tooth of S.
meltoni (72 degrees); all comparisons were made with specimens
which exhibited little wear and are known or inferred to be mis
based on a complete metalophid. Consequently, the trigonid is
relatively wide in comparison with the talonid, falling near the
limit of variation exhibited by small Blancan ground squirrels
(Fig. 3). The talonid basin lacks rugosity and slopes gently downward towards the ectolophid. The entoconid is bounded mesially
by a shallow notch. The posterolophid forms a slight arc from the
entoconid to hypoconid, and the ectolophid, connecting hypoconid with protoconid, lacks any evidence of a mesoconid. The hypoconid projects buccally and lacks strong mesial deflection.
The buccal valley, viewed buccally, is broadly U-shaped with
a flattened floor (Fig. 4B), differing from typical specimens of S.
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MARTINET AL.-BLANCAN RODENTS AND LAGOMORPHSFROM KANSAS
1077
The hypoconidextends mesiobuccally,with a more mesial derowly U-shapedor V-shapedprofile.The lingualmargin,viewed flectionthanis typicalof Spermophilussp. (small)fromDeerPark
lingually,mergesgraduallywith the distalface of the metaconid. B and resemblingtypical specimensof S. rexroadensisand the
Taxonomicassignmentof this specimenis problematical.
Sper- type of S. finlayensis.In buccal view, the buccal valley of both
mophilus meadensis, S. howelli, and S. meltoni are all similar teeth is broadenedbasally,resemblingtypicalspecimensof S. cf.
dentally;furtherwork is needed to clarify relationshipsamong rexroadensisbut unlike the type and single specimenof S. finthese small Blancansquirrels.The broadlyU-shapedbuccal val- layensis which exhibits a compressed,V-shapedbuccal valley.
ley and relativelynarrowtalonidcomparedto trigonidare unlike However, this character varies within S. rexroadensis. Both teeth
or at the limit of variationobservedin these knownBlancanspe- fall within metric variation of S. cf. rexroadensis from Fox Cancies. Additionalmaterialis neededto determinewhetherthis rep- yon and near S. finlayensis from Hudspeth but outside variation
resentsa new species and to clarifyaffinitieswith describedspe- in the sample of S. rexroadensis from Rexroad Loc. 3 (Fig. 3).
The single m3 (VP-14315) is extremely worn and eroded, but
cies.
meadensis, S. howelli, and S. meltoni which display either a nar-
appears to retain a trace of a talonid trench lingual to the ectoSPERMOPHILUS
(Hibbard, 1941)
cf. REXROADENSIS
lophid and a basally broad buccal valley.
Figure 4.3, 4.4
The specimens from Deer Park B most closely resemble S. cf.
Material examined.-FHSM VP-14224 R ml or m2; 14239, R rexroadensis from Fox Canyon, which is early Blancan in age,
p4; 14240, L ml or m2; 14241, L M1 or M2; 14242, R M1 or than any other Blancan squirrel known from the Great Plains. The
M2; 14243, L M3; 14244, R ml or m2; 14315, R m3.
relationships among the Fox Canyon squirrel, S. rexroadensis
Discussion.-Two upper cheek teeth probably represent RM1 from the younger Rexroad Locality 3 1.f., and the late Blancan S.
(VP-14242; L = 2.76 mm, W = 3.40 mm) and LM2 (VP-14241; finlayensis remain unclear; they may represent the same lineage,
L = 2.64 mm, W = 3.25 mm). The anterior cingulum of M1 although additional material is needed for S. finlayensis. Given
tapers lingually and merges gradually with the mesial surface of this uncertainty, the occurrence of S. cf. rexroadensis from Deer
the protocone in occlusal view (Fig. 4.4). In lingual or buccal Park B provides little biochronologic information.
view, the protocone is high, bears a steep mesial face, and contacts the anterior cingulum at an abrupt angle (Fig. 4.3). The
Gill, 1872
Family GEOMYIDAE
protoloph is high and complete; the metaloph is sharply constrictGEOMYS (GEOMYS) cf. QUINNIMcGrew, 1944
ed at its contact with the protocone and bears a prominently swolFigure 2.4-2.13
len metaconule (Fig. 4.4). The central valley between protoloph
Material
examined.-FHSM
VP-14195, eight p4s; 14424, L
and metaloph is narrow and partially blocked buccally by a promR
14426-27
L
14428 L P4; 14196, seven
14425,
p4;
p4;
p4s;
inent mesostyle closely appressed to the metacone (Fig. 4.3). In
25 upper incisor pieces; 14198, five P4s; 14199, eight
P4s;
14197,
of
the
anterior
the
M2
does
not
contrast,
cingulum
putative
taper
as strongly towards its lingual terminus and deflects distally to upper incisor pieces; 14200, deciduous p4; 14283, part edentulous
contact the protocone. In lingual view, the mesial surface of the palate; 14284, part edentulous L mandible; 14285, 55 molar and
incisor pieces.
protocone merges gradually with the anterior cingulum. The meDiscussion.-Two species of pocket gopher are present in the
is
at
contact
with
the
constricted
its
the
but
taloph
protocone,
metaconule is less developed than on Ml. The metaloph angles Deer Park A and B material. The smaller taxon will be considered
below. There are two robustBlancanspecies of Geomysrecogaway from the protoloph as it courses buccodistally, resulting in
a buccally broadened central valley. There is no mesostyle. The nized from the Meade Basin record.The first is Geomysquinni,
Ml closely resembles some specimens of S. rexroadensis from named by McGrew (1944) from the Sand Draw 1.f. Hibbard
Rexroad Loc. 3 and S. cf. rexroadensis from Fox Canyon; upper (1956, 1967) identified this species from Deer Park A. The second
teeth are unknown from S. finlayensis. The buccally expanded is G. jacobi, named by Hibbard (1967) and including material
central valley as seen on the putative M2 is present in at least from Rexroad Loc. 3 and Wendell Fox Pasture (loc. UM-K3-53).
some specimens from Rexroad Loc. 3. The single M3 is heavily Hibbard and Riggs (1949) also reported G. jacobi from the Keefe
worn, chemically eroded, and preserves little morphological in- Canyon 1.f., but the pocket gopher material from that fauna may
include two species, neither of which is G. jacobi (R. Martin,
formation.
The single p4 is too heavily eroded to measure but exhibits a personal observation). According to Hibbard (1967), G. jacobi is
relatively narrow trigonid compared to the talonid, a distally open separable from G. quinni on the basis of its smaller size and less
trigonid pit between protoconid and metaconid, and a distinct developed retromolar fossa lateral to m3 (terminology follows
trenchon the talonidfloor adjacentto the ectolophid.The basic Wilkins, 1984). There are no skulls or complete mandibles from
morphologyresemblesspecimensof S. rexroadensis,althoughthe Deer Park, and we have not recovered any from the numerous
new localities we have discovered. It seems obvious that we will
lattertypicallydisplaya less-developedtalonidtrench.
Threespecimensrepresentml or m2, but one is so worn (VP- have to develop a replacement chronology and character analysis
14244) as to providelittle information.The two remainingspec- for Meade Basin gophers that is based primarily on the interplay
imens differ in size and morphology. The smaller (VP-14240; L of dental features and, in some cases, characters of the anterior
= 2.46 mm, TrW = 2.68 mm, TIW = 3.01 mm) exhibits a disportion of the mandible, the latter of which is occasionally reptinct trigonid pit and relatively narrow trigonid compared to the resented.
One of us (RAM) recently compared the large Deer Park A
talonid, and probably represents ml. The larger specimen (VP14224; L = 2.71 mm, TrW = 3.09 mm, T1W = 3.17 mm) is and B Geomys dental and mandibular material with G. quinni
subequal in width across the trigonid and talonid and bears a material from Sand Draw in the University of Michigan collectrigonid pit, but this pit is confluent with a shallow groove which tions, and could detect no significant qualitative differences. There
descends to the floor of the talonid. On both specimens, the tal- is considerable overlap for most dental dimensions (Table 1), alonid is bounded distally by a continuous, arcing posterolophid though the Sand Draw specimens appear to have, on average,
that courses from an indistinct entoconid to the hypoconid, and slightly longer and wider p4s. On this basis, and on the absence
buccally by the ectolophid which bears no trace of a mesoconid. of observed qualitative features, we tentatively accept Hibbard's
A shallow trench lies lingual to the ectolophid on both teeth, (1956) identification and refer the material from Deer Park A and
resembling S. finlayensis and many specimens of S. rexroadensis. B to G. quinni. Table 1 also shows there is much overlap between
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1078
JOURNAL OF PALEONTOLOGY,V. 76, NO. 6, 2002
the Sand Draw and Deer ParkG. quinniand samplesfrom Rexroad Loc. 3 and WendellFox Pasturereferredto G. jacobi. Althoughthese datasuggestthatthe MeadeBasingopherfromRexroad Loc. 3 and WendellFox Pasturemay be the same as the
Deer Parkspecies,we will refrainfroma formalrecommendation
until studiesunderwayarecompleted.Some p4s are illustratedin
Figure2. Note the U-shapedreentrantsfilled with cement.In Pliogeomys buisi (Hibbard,1954a) from Buis Ranch, these reentrantstend to be V-shaped,and cementis lacking.
GEOMYS(NERTEROGEOMYS)
cf. MINORGidley, 1922
Figure 2.14-2.17
Material examined.-FHSM VP-14201, one p4 and one P4;
14202, four upper incisor pieces; 14203, misc. molars; 14204,
fourP4s; 14205,threep4s; 14206, 15 upperincisorpieces; 14429,
R p4; 14430 L p4.
Discussion.-Teeth of a small pocket gopherfrom Deer Park
morphologicallymatchthose referredto Geomys(N.) minorfrom
the Rexroad Loc. 3 l.f. (Hibbard, 1967). However, there is an
obvious size difference between the teeth from Deer Park B and
those referred to G. minor from, for example, Rexroad Loc. 3
(Table 1, Fig. 2). Without mandibularmaterial of the small gopher
from the Deer Park localities we cannot be certain of the species
identification,and we only tentativelyrefer the materialto G.
minor.
Allen and Chapman, 1893
Family HETEROMYIDAE
PEROGNATHUS
cf. GIDLEYI
Hibbard, 1941
Material examined.-FHSM VP-14286, R P4; 14287, R M1;
14288, R M3; 14289, L M3; 14290, L p4; 14291, R ml.
Discussion.-The RP4 protolophlacks accessorycusps, con-
sisting only of the protocone. The metaloph cusps are connected
by a broad dentine lake and are no longer individually distinguishable. At this stage of wear the protoloph and metaloph are
joined by a broad dentine connection that is slightly labial of the
midline of the tooth. The lingual part of the transverse valley is
slightly deeper than the labial part. The P4 is three-rooted.
The RM1 is bilophodont, with the metaloph slightly narrower
transversely than the protoloph. Neither an anterior cingulum nor
a small style is present on the anterior wall of the protoloph. Wear
has united the paracone and protocone, and the enamel wall separating the protostyle and the protocone is starting to be breached.
The protoloph is widest at the protocone. On the metaloph the
metacone, hypocone, and hypostyle are all united, and the hypostyle forms a pronounced anterolingual inflection. The metaloph is widest at the hypocone. The transverse valley is deepest
labially, and with greater wear the lophs will unite first on the
lingual side. The protostyle and hypostyle are well separated and
not strongly convergent as they are in M2, and VP-14287 is not
as anteroposteriorly compressed as M2, both of which features
help identify this tooth as Ml rather than M2. The roots are broken at their junction with the crown.
The M3 is subcircular with the protoloph wider than the metaloph. In both Deer Park specimens the individual cusps are no
longer distinguishable due to wear. Wear has closed off the transverse valley on the lingual side in both specimens; the labial part
of the transverse valley is still open. Two separated roots are
presenton VP-14289.
On the Lp4 the metalophidis much narrowerthan the hypolophid. Wear has proceeded to the stage where individual cusps
are no longer distinct, the metalophid and hypolophid are united
centrally, and the anterior reentrant valley between the protoconid-metaconid complex and the protostylid is eliminated. There is
no hint of an accessory cusp, or anteroconid, on the anterior wall
of the metalophid. The labial reentrantvalley is slightly shallower
than the lingual reentrantvalley and does not extend as far toward
TABLE2-Measurements of Perognathus cf. gidleyi from Deer Park B. GL
= greatest length in side view. TrWAn = greatest transversewidth of the
anterior lobe in occlusal view. TrWPo = greatest transversewidth of the
posterior lobe in occlusal view.
FHSMno.
14286
14287
14288
14289
14290
14291
Tooth
GL
RP4
RM1
RM3
LM3
Lp4
Rml
1.23
1.08
0.73
0.67
0.83
1.05
TrWAn
0.69
1.15
0.87
0.80
0.63
1.06
TrWPo
1.30
1.13
0.70
0.67
0.80
1.07
the base of the crown. At this stage of wear there is no indication
of a posterior median valley on the back wall of the hypolophid,
if it were ever present.
On the Rml the metalophid and hypolophid are nearly the same
width with three cusps each. A narrow, posteriorly flexed, anterolabial cingulid extends from the protoconid to the protostylid.
The metaconid and protoconid show only apical wear, and are not
yet connected by a dentine bridge. On the hypolophid, the hypoconid is the largest cusp and the hypostylid is the smallest and
lowest of the three cusps. The entoconid and hypoconid also show
apical wear only. The transverse valley is flexed slightly posteriorly on either side of the hypoconid, is shallowest centrally, and
is slightly deeper lingually than labially, thus in extreme wear the
metalophid and hypolophid will unite sooner labially than lingually. The fused anteriorroots are preserved nearly to their distal
ends; the posterior roots are broken off just below the crown.
There appears to be only one species of Perognathus represented at Deer Park B. The six teeth are closest in size (Table 2)
and morphology to the teeth of P. gidleyi (Hibbard, 1941b) from
Rexroad Loc. 3. The Deer Park B Perognathus is larger than P.
pearlettensis (Hibbard, 1941a) from early Blancan through early
Irvingtoniandepositsof the Meade Basin, P. henryredfieldi(Jacobs, 1977) of the HemphillianRedingtonl.f. of Arizona,andP.
dunklei (Hibbard, 1939) from the Hemphillian Edson quarry of
Kansas, and slightly smaller than P. rexroadensis (Hibbard, 1950)
from Fox Canyon, except for the ml (VP-14291), which is similar
in size to that of P. rexroadensis (cast of UMMP 28248). In
addition, the Deer Park Perognathus differs from P. mclaughlini
in lacking the prominent, low, anterior cingulum on Ml, present
on specimens from Arizona (Jacobs, 1977), and in having a deeper labial reentrantvalley which is only slightly shallower than the
lingual reentrant valley on the p4, resulting in a central union of
the metalophid and hypolophid. In P. mclaughlini the labial valley
is much shallower than the lingual valley, such that the union of
the metalophid and hypolophid occurs first on the labial side rather than centrally (Hibbard, 1949; Jacobs, 1977). Labial, rather
than central, union of the p4 lophids also characterizes P. henryredfieldi (Jacobs, 1977). The labial reentrant valley is also notably shallower than the lingual valley on the p4 of P. rexroadensis (Hibbard, 1950), a feature that contrasts with P. gidleyi. On
the Deer Park B p4, the lingual valley is only slightly deeper than
the labial valley, with the difference in depth less than seen in P.
rexroadensis (cast of UMMP 28248). The Deer Park B p4 also
lacks the hypostylid seen on the type of P. maldei from the late
Blancan Hagerman 1.f. of Idaho (Zakrzewski, 1969). The worn
Deer Park B Ml differs from that of P. stevei from the Hemphillian Ordnance locality of Oregon in lacking an anterior cingulum from the paracone or protocone to the parastyle. J. Martin
(1984) reported a poorly formed anteriorcingulum on P. rexroadensis, and Albright (1999) reported the presence of a small style,
rather than an anterior cingulum, anterior to and between the Ml
paracone and protocone on P. rexroadensis from Fox Canyon, a
feature absent on VP-14287 from Deer Park B. In P. gidleyi the
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MARTINET AL.-BLANCAN RODENTS AND LAGOMORPHSFROM KANSAS
1079
Ml anteriorcingulumreportedlyvariesfrom continuousto weak TABLE3-Measurements of Prodipodomysfrom Deer ParkB. GL = greatest
length in side view. TrW = greatest transverse width in occlusal view.
anddiscontinuous(Tomida,1987).Perognathusstrigipredusfrom
LabHt = Labial dentine tractheight. LinHt = Lingual dentine tractheight.
the early BlancanHouse Mt. Loc. 318 of the VerdeFm. of Arizona (Czaplewski,1990) differsfromthe Deer ParkB specimens
FHSM
in having a relativelywider metalophidon the p4 that is nearly
No.
Tooth
GL
TrW LabHt LinHt
Species
as wide as the hypolophid,deeperlabialandlingualreentrantson
14292
RM2?
1.00
1.27
0.00
0.00
Prodipodomyssp.
p4, and ml that is anteroposteriorlycompressed and higher cf. P. idahoensis
14293
RM2
1.00
1.37
0.38
0.32
crowned.
14294
Lm3
0.76
1.03
0.17
The Deer Park B specimens seem closest in size and morphology to P. gidleyi; a possible differenceis the lack of an anteriorcingulumon the wornDeer ParkMl, althoughthe anterior Edsonlocalityto be nearlyfused.Measurements
of the Deer Park
cingulumon P. gidleyi is variablein strength(Tomida,1987). In B Prodipodomysare presentedin Table3.
Kansas,P. gidleyi has been reportedfrom the RexroadLoc. 3
PRODIPODOMYS sp.
and Nash 72 1.f.s (Martinet al., 2000).
PRODIPODOMYScf. IDAHOENSISHibbard,1962
Material.-FHSM VP-14293, R M2; 14294, L m3.
Comments.-VP-14293 is recognized as an M2, having the
protolophnoticeablywider than the metaloph.The tooth is well
wornwith the individualcuspsof each lobe indistinguishable.
The
transversevalley is still open labially,withthe reentrantextending
about 35 percentacross the width of the occlusal surfaceat the
protoloph.On the labial surfacethe reentrantbetweenthe protoloph and metalophgraduallylessens in depthtowardthe base of
the enamel,anddisappearsjust abovethe top of the dentinetract.
Distinct labial and lingual dentinetractsare present.The labial
dentinetractis bilobed,with the lobe below the protolophtaller
than the lobe below the metaloph.The lingual dentine tract is
wedge shapedwith a single,pointedapex.Threerootsarepresent
which are fused except for theirdistalends.
The m3 is well worn with the individuallophidcusps obliterated. The crown is roughlyoval, with the anteriorenamel wall
(metalophid)slightly flattertransversely,wider,and higherthan
the posteriorly-convexposteriorenamel wall (hypolophid).The
presumedlingualmarginof the occlusal surfaceis broader(less
pointed)thanthe labial margin.Only a single root is present.An
incipientdentinetractis presenton the lingualside of the tooth.
The length, width and dentinetractheightsfor VP-14293 fall
within the range of measurementsfor the M2 of P. idahoensis
from the Verdel.f. (Czaplewski,1990), being very close to the
means for anteroposterior
length and labial dentinetractheight.
VP-14293 however,is slightly longer,narrower,and has shorter
dentinetractsthanthe isolatedM2 of P. idahoensis(UCR22460)
fromthe San TimoteoFormation,California(Albright,1999),and
is the same lengthbut slightly narrowerthanP. idahoensisfrom
the Yep6mera1.f.,Mexico (LindsayandJacobs,1985). Both VP14293 andthe Yep6meraM2 have rootsthatare separatedistally,
and UCR 22460 is describedas havingwell-developedbut fused
roots (Albright,1999). The m3 from Deer Park B is the same
size and very similar morphologicallyto P. idahoensis (IGCU
2717:cast) from the Yep6mera1.f.; the dentine tract for IGCU
2717 (illustratedby LindsayandJacobs(1985)) is the sameshape
andheightas thatof the Deer ParkB specimen.VP-14293is also
similarin size to P. idahoensisfrom the Verde1.f. (Czaplewski,
1990),but is slightlynarrowerandhas shorterdentinetracts.Prodipodomystiheniis reportedto have less fused roots and shorter
dentinetractsthanP. idahoensis(Czaplewski,1990). Althoughit
Material examined.-FHSM VP-14292, R M2?
Discussion.-This toothis eitherMl or M2, but morelikely to
be M2 based on overallproportions.The individualcusps of the
protolophandmetalophareobliteratedandmergedthroughwear.
The protolophis transverselywiderthanthe metaloph.The transverse valley is still open labiallyand extendsroughly40 percent
across the width of the occlusal surface.On the labial surface,
the valley between the protolophand metalophends 0.43 mm
above the base of the enamel. Dentine tractsare absent.Three
widely separatedroots are present,a largeroot linguallyandtwo
roots labially.
VP-14292 is assignedto the genus Prodipodomysratherthan
to Dipodomysbecause of the absence of dentinetractsand the
presenceof threeseparateroots. In Prodipodomys,dentinetracts
areabsentor only slightlydevelopedandmultiple,divergentroots
are presentin the cheek teeth (Zakrzewski,1981; Dalquestand
Carpenter,1986;Dalquestet al., 1992).VP-14292differsfromP.
idahoensisin being transverselynarrowerand lackingthe incipient dentinetractsfoundin thatspecies (Zakrzewski,1969;Lindsay and Jacobs, 1985; Czaplewski,1990; Albright, 1999). VP14292 is slightly longer but the same width as P. kansensisreportedfrom the Redingtonl.f., Arizona;the roots of the Redington specimen differ from those of VP-14292 in being fused
proximallyand weakly separateddistally(Jacobs,1977). An M2
from Edson Quarryquestionablyreferredto P. kansensis(Hibbard,1939) is similarin size to VP-14292,with a reportedtransverse widthof 1.2 mm and a length (takenfrom the illustration)
of about1.05 mm;however,the roots aresaid to be nearlyfused.
VP-14292 almostexactly matchesin size the M2 of P. centralis
F:AM 87427 from the Keim Fm. of Nebraska(Hibbard,1972).
The teeth of P. centralis are said to lack dentinetractson the
sides of the teeth at the base (Hibbard,1954b, 1972).
Based on the publisheddescriptionsof P. griggsorumand P.
tiheni,VP-14292cannotbe distinguishedfromthesetwo species.
AlthoughP. tiheniis sometimessaid to have the highestdentine
tractsof the species of Prodipodomys(Zakrzewski,1981), Czaplewski(1990) reportedthe dentinetractsin P. tihenito be shorter
thanin P. idahoensis;he describedthe dentinetractsof P. tiheni
as broad,roundedemarginationsat the bases of roots.
FamilyCRICETIDAERochebrune,1883
Subfamily SIGMODONTINAE
Wagner, 1843
PEROMYSCUS
cf. HAGERMANENSISHibbard, 1962
Figure5
Material examined.-FHSM VP-14149 L ml; 14150, R ml;
differs in having three well-separatedM2 roots and in lacking 14151, L m2; 14152, L Ml; 14153, R ml; 14154, R M2; 14156,
dentinetracts(Hibbard,1954b;1972). Prodipodomysgriggsorum threeR Mls, L Ml; 14157, L M2; 14159, R ml; 14368, L M1;
also has three well-developed roots on the M2 (Zakrzewski, 14405-6, two R mls.
Discussion.-Teeth of Peromyscus from Deer Park B are quite
1970). The M2 of P. kansensisfrom the Redington1.f.is slightly
smallerthan VP-14293, but is similarin having the roots fused bunodontwith strongcusps andrelativelythickenamel.The Mis
proximallyand weakly separateddistally(Jacobs,1977);Hibbard show a bilobedanteroconewith an anteromedian
groove,even in
(1939) also reportedthe roots of referredP. kansensisfrom the worn specimens.The anterolabialstyle is stronglydevelopedand
is the same size as VP-14293, P. centralis (=P. rexroadensis)
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1080
JOURNAL OF PALEONTOLOGY,V. 76, NO. 6, 2002
3
2:
12
45
6
10
4
11-
7
8
I0
3
6
FIGURE
6-Baiomys rexroadiHibard,1941 from Deer ParkB. 1, R ml
(FHSMVP-14185);2, L ml (FHSM 14186);3, posteriorfragmentof
R ml (FHSMVP-14187);4, L m2 (FHSMVP-14191);5, Rm3 (FHSM
VP-14356);6, RM2 (FHSMVP-14355).Scale bar = 1.0 mm.
9
1'-
FIGURE
5-Peromyscus cf. hagermanensisfrom Deer ParkB. 1, L Ml
(FHSMVP-14152);2, R Ml (FHSMVP-14405);3, L Ml (FHSMVP14368);4, L M2 (FHSMVP-14157);5, R ml (FHSMVP-14150);6,
L ml (FHSM VP-14149); 7, R ml (FHSM VP-14406); 8, R ml
(FHSM VP-14159);9, R ml (FHSM VP-14188); 10, R m2 (FHSM
VP-14154);11, L m2 (FHSMVP-14151).Scale bar = 1.0 mm.
attached to the labial side of the anterocone in worn specimens.
The mesoloph on Ml and M2, extended from the anterior arm of
the hypocone, varies from absent to well developed. The mesoloph does not reach the mesostyle in any specimen. One of the
M2s has a double connection between paracone and protocone.
The mls have a double anteroconid, although it is only visible in
lightly worn specimens. The metalophulid is transverse. The lingual anterior cingulum is strong in mls and m2s. The ml and m2
show a cingular mesostylid of variable development. Half of the
mls have an anterior spur in the entoconid directed towards the
mesostylid.
The teeth from Deer Park B are similar in size (Table 4) to P.
hagermanensis from the Hagerman l.f. (Zakrzewski, 1969). The
morphology, although variable, is also similar in both assemblages. The Peromyscus specimens from Deer Park B are considerably
larger than the material of P. baumgartneri from Rexroad Loc. 3
(Hibbard, 1954b) and P. cragini from the Cudahy l.f. (Hibbard,
1944). The latter species also have very simple molars, without
accessory structures (Hibbard, 1954b; Paulson, 1961). Peromyscus kansasensis from Rexroad Loc. 3 is a larger species, but also
has an ml without an anteromedian groove and lacks accessory
structures on all molars (Hibbard, 1941b). Peromyscus sawrockensis (Hibbard, 1964) was recently reassigned to the genus Symmetrodontomys (Martin, 2000). Peromyscus nosher Gustafson
(1978) from the White Bluffs l.f. of Washington is similar in size
and morphology to P. hagermanensis. These species differ in the
more distinctly bilobed anteroconid on the ml of P. nosher (Albright, 1999) and in the small and cuspoid parastyle of P. nosher
compared to the large and lophoid parastyle in P. hagermanensis
(Gustafson, 1978; Tomida, 1987).
BAIOMYSREXROADI
Hibbard, 1941b
Figure 6
Material examined.-FHSM VP-14185, R ml; 14186, L ml;
14187, R ml; 14189, R ml; 14190 L M1; 14191 L m2; 14245,
R ml; 14354 R ml; 14355 R M2; 14356 R m3.
Discussion.-Baiomys material from Deer Park B show close
similarities in size (Table 5) and morphology to the type material
of B. rexroadi from Rexroad Loc. 2a and is therefore assigned to
this species. Baiomys sawrockensis (Hibbard, 1952a) from the
Saw Rock Canyon 1.f. has mis that are slightly shorter because
of a less developed procingulum (sensu Hershkovitz, 1962).
Baiomys kolbi from Fox Canyon (Hibbard, 1952b) supposedly
differs from B. rexroadi from the same locality by virtue of its
simple, rather than bifid, anteroconid on ml. Size of the two species is very similar, and this species pair needs to be re-evaluated
in light of current knowledge of population variation.
Baiomys aquilonius Zakrzewski (1969) has a peculiar ml with
a distinct anterostylid that confers an expanded shape to the procingulum. The m3 of this species is smaller than that of B. rexroadi from Rexroad Loc. 2a and Deer Park B.
SYMMETRODONTOMYS
SIMPLICIDENSHibbard, 1941b
Figure 7
Material examined.-FHSM VP-14155 R m3; 14160, R ml;
14161, R m2, L m2; 14162, L m3, R m3; 14163, two L Mls,
two R Ml; 14166, RM1; 14167, R m2; 14168, L m3; 14169,
three R mls; 14170, R M2, L M2; 14246, LM1; 14372 R m2;
14373 R m2; 14374 L M3; 14375 R M2; 14407-8 two R m2s;
14409 L m3; 14410, R m3; 14412, R Ml; 14413 L Ml; 14414
L m2; 14415 L m3; 14416, R ml; 14417 L ml; 14418-9 two R
M2s.
TABLE
molarsfrom
4-Length andwidthof Peromyscuscf. hagermanensis
DeerParkB. N = numberof specimens,Min. = smallestmeasurement, TABLE
5-Length andwidthof BaiomysrexroadimolarsfromDeerParkB.
Max. = largestmeasurement,
s.d. = standard
deviation.
as in Table4.
Abbreviations
ml
m2
M1
M2
N
Length
Min. Mean Max.
5
2
3
2
1.51
1.34
1.72
1.30
1.60
1.81
1.66
1.34
1.89
1.43
Width
s.d.
N
Min. Mean Max.
s.d.
0.06
7
2
3
2
0.97
0.99
1.09
0.02
0.09
1.01
1.11
1.04
1.03
1.15
1.10
0.03
N
ml
m2
m3
M2
4
1
1
1
Width
Length
Min. Mean Max.
s.d.
N
Min. Mean Max.
s.d.
1.18
0.04
4
1
1
1
0.74
0.03
1.23
0.89
0.69
0.95
1.28
-
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0.77
0.72
0.56
0.73
0.80
MARTINET AL.-BLANCAN RODENTS AND LAGOMORPHSFROM KANSAS
1081
TABLE6-Measurements of Symmetrodontomyssimplicidens molars from
Deer Park B. Abbreviationsas in Table 4.
:t.
1
I
2
3
ml
m2
m3
M1
M2
i-~~ I
4
,
5
6
12 1213
c
14
10
Length
Min. Mean Max.
1
4
5
3
4
1.69
1.35
1.13
1.81
1.28
M31
7
g
9
N
11
15
FIGURE
7-SymmetrodontomyssimplicidensfromDeer ParkB. 1, L Ml
(FHSMVP-14412);2, R M1 (FHSMVP-14413);3, L M2 (FHSMVP14170);4, R M2 (FHSMVP-14418);5, R M2 (FHSMVP-14419);6,
R M2 (FHSM VP-14375); 7, L M3 (FHSM VP-14374); 8, R ml
(FHSMVP-14416);9, anteriorhalf of L ml (FHSMVP-14417);10,
R m2 (FHSM VP-14407); 11, R m2 (FHSM VP-14408); 12, Lm2
(FHSMVP-14414);13, L m3 (FHSM 14409); 14, R m3 (FHSMVP14410); 15, L m2 (FHSMVP-14415).Scale bar = 1.0 mm.
Discussion.-The upper molars show buccal and lingual cusps
aligned. Almost all Mis and M2s have a mesostyle. The mesostyle is almost always connected to a mesoloph. The latter structure runs from the posterior wall of the paracone towards the
mesostyle. In Ml there is also a well-developed parastyle.
The lower molars have a short mesolophid that runs from the
anterior wall of the entoconid. One ml displays a mesostylid. The
lingual and buccal cusps are slightly more alternate than in the
upper molars.
Some very worn m3s have been tentatively assigned to this
species based on size and general pattern; they could belong to
Peromyscus or Onychomys. Measurements are given in Table 6.
BENSONOMYS
Hibbard, 1956
MEADENSIS
Figure 8
Material examined.-FHSM VP-14158, R MI; 14165 2 L
M2s;14184 R ml; 14247, L Ml; 14248, R ml; 14249, L ml;
14250, R ml; 14251, R ml; 14252, L m2; 14357, L m2; 14358
L m2; 14359 R m2; 14360 L M2; 14361 R M2; 14362 L M2.
Discussion.-Most of the specimens are described individually
below, followed by comparisons with related forms.
VP-14247: L Ml. The major cusps are alternate. There is a
tubercle below the paracone, a vestigial fourth root. The anterocone is transversely broad and divided into two lobes by a shallow, anteromedian groove. The labial lobe is larger than the lingual lobe. A low cingulum with two styles is present at the anterior base of the anterocone. The anterior arm of the protocone
connects with the labial lobe. A low cuspule is on the labial border, between the anterocone and paracone. The posterior cingulum
connects the hypocone with the metacone.
VP-14158: R Ml. The roots of this specimen are not preserved.
1.32
1.06
1.73
1.19
1.37
1.26
1.95
1.41
0.86
Width
s.d.
N
Min. Mean Max.
s.d.
0.02
0.08
0.12
0.10
2
4
5
5
4
1.13
1.11
1.05 1.08 1.13
0.94 0.97 0.99
1.08 1.17 1.26
1.09 1.14 1.19
0.03
0.02
0.07
0.04
-
1
0.91
Relative to 14247, 14158 lacks the low cingulum at the anterior
base of the anterocone.
VP-14360: L M2. There is a poorly-developed anterior cingulum that delimits a shallow and broad protoflexus. The main cusps
are almost aligned transversally. The hypoflexus is curved anteriorly and the lingual end is closed by a low cingulum that runs
between the bases of the protocone and hypocone. The metacone
connects to the posteroloph just before the latter ends. The paracone shows a small posterior spur. The remaining M2s (VP14361, 14362, 14165) are badly preserved and heavily worn.
Nevertheless, the main morphological characteristics pointed out
in VP-14360 can be observed on them.
VP-14249: L ml. The anteroconid is not broad, forming the
apex of a triangle anteriorly. It is bilobate, divided by a moderately deep anteromedian groove. The external and internal lobes
are subequal and broadly confluent posteriorly. There is a low
anteromedian stylid attached mainly to the labial lobe of the anteroconid. The anterolophid is short and is connected medially to
the metalophid/anterior arm of the protoconid. The anterior cingulid extends along the labial margin from the base of the protoconid anteriorly and terminates in a small stylid which is separated from the base of the labial anteroconid by a narrow notch.
The valley (metaflexid) between the lingual anteroconid and
metaconid is narrow. The metalophid extends backward at about
/0-b
0
11
fA
mom
S3
/2
4
5
FIGURE
8-Bensonomys meadensisHibbard,1956 fromDeer ParkB. 1,
R Ml (FHSM VP-14158); 2, L M2 (FHSM VP-14360); 3, R ml
(FHSM VP-14184); 4, L m2 (FHSM VP-14357); 5, R m2 (FHSM VP14358). Scale bar = 1.0 mm.
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1082
JOURNAL OF PALEONTOLOGY,V. 76, NO. 6, 2002
TABLE7-Measurements of Bensonomysmeadensis molars from Deer Park
B. Abbreviationsas in Table 4.
this locality (contraCzaplewski,1987a).Hibbard(1972) laterreferred specimensfrom the Sand Draw 1.f. of Nebraskato this
species. Hibbard(1956) distinguishedB. meadensisfromB. ariWidth
Length
zonae, then only known from the Blancan(ca. 3.2 Ma) Benson
N Min. Mean Max. s.d.
N Min. Mean Max. s.d.
fauna of Arizona,by the more dorsal and posteriorposition of
the massetericcrest. Hibbard(1956) also describedB. eliasi,
ml
3 1.55 1.58 1.60 0.03
3 0.95 0.98 1.02 0.04
m2
2 1.23
1.24
3 0.98 1.04 1.09 0.06
which is fromRexroadLocs. 2 (holotypelocality)and 3 andFox
2 1.62
2 0.99
1.65
1.02
M1
Canyon.The holotype of B. eliasi, initially describedas PeroM2
3 1.12 1.14 1.18 0.04
3 0.88 0.92 0.96 0.04
myscuseliasi (Hibbard,1938), is a mandiblewith heavily worn
ml-m3. Bensonomyseliasi was distinguishedfrom the similar
sized B. arizonaein havinga widerlower incisorand an ml that
30 degreesfrom its juncturewith the anteriorarm of the proto- is more rectangular,with a broadversus triangularanteroconid,
conid towardthe metaconid.The apex of the metaconidis just andbroaderreentrantvalleys (Hibbard,1956). A relativelybroad
anteriorto the apex of the protoconid.The entoconidis opposite anteroconidis presentin most of nine specimensfromFox Canthe hypoflexid.Thereis a minuteectostylidon the very low labial yon andone fromRexroadLoc. 3 (Hibbard,1950,pl. I), although
cingulumbetweenthe protoconidand the hypoconid,whichcon- some, particularlythe smallerspecimens,have narrowanterocontinues aroundonto the posteriormarginof the hypoconid.The ids. Hibbard(1950) speculatedthatmorethanone species might
posteriorcingulum extends posterolinguallyfrom the posterior be present.Bensonomysmeadensis was distinguishedfrom B.
armof the hypoconidand then transverselyso thatit is close to, eliasi by its smallersize (ml-m3 length 3.5 vs. 3.8 mm) and
but not in contactwith, the entoconid.
narrowerlowerincisor.HibbardconsideredbothB. meadensisand
VP-14184:R ml. Similarto VP-14249in size andmorphology, B. arizonae to have similarly shaped ml's, with a triangularit differs by having a more stronglyanteromediallydivided an- shapedanteroconid,unlikethe broadanteroconidhe attributedto
teroconid.The anterolophiddoes not reachthe metalophid/anter- B. eliasi. Czaplewski(1987a) statedthat in the absenceof quanior armof the protoconid.The ectostylidis strongerthanin VP- tified differencesand comparativespecimenshe was unableto
14249. The posteriorcingulumshows a well-developedpostero- distinguishspecimensfrom the early Blancanpart of the Verde
conulid.
Formationof Arizonathat he referredto Calomys(Bensonomys)
VP-14248: a slightly worn, R ml. The labial anteroconidis arizonae from B. eliasi or B. meadensis. Czaplewski (1987b) also
brokenoff andtheredoes not appearto have been an anterostylid. referredthe SandersB. meadensisandspecimensfromthe middle
Relative to VP-14249, the anteroconidappearsto have been and late Blancanof Arizonato C. (B.) arizonae.Additionalspecbroadertransverselyand more stronglydivided anteromedially. imensfromthe RexroadFormation,collectedfor theMeadeBasin
The metaflexidis wider.
RodentProject(Martinet al., 2000), tentativelysupportthe recVP-14250:anteriorhalf of R ml. The tooth is very worn. An ognitionof B. eliasi as a distinctspecies.
anterolophextendsfromthe bilobedanteroconidto the protoconONYCHOMYS
Carleton and Eshelman, 1979
cf. HOLLISTERI
id.
Figure9
VP-14251:anteriorhalf R ml. The bilobed anteroconidis diMaterial examined.-FHSM VP-14164, M2; 14209, R ml;
vided by a stronganteromediangroove.
VP-14252:L m2. The toothis brokenanteriorto the protoconid 14210, R m2; 14211, L m2; 14212, R M1; 14213, L m2, R m2;
and metaconid.The cusps are alternate.The posteriorcingulum 14214, two L mls, R ml; 14215, R M2; 14363, RMI; 14364, L
is short, terminatingbefore reachingthe lingual marginof the m3; 14365, L m3; 14366, Lm3; 14367, R m3; 14411, R M2;
tooth.
14431-33, threeR Mls; 14434, L m2; 14435, R m2; 14436, L
Althoughwe recognize that Bensonomysis a primitivephyl- m2; 14437, L ml.
Discussion.-Teeth of Onychomys from Deer Park B have a
lotineclosely relatedto Calomys(Baskin,1978, 1979, 1986;Martin, 2000), becauseof its plesiomorphicfeatureswe returnBen- generalpatternof relativelysectorialandslendercusps.The cusps
alternate,especially in the lower molars.The anteroconeof the
sonomysto genericstatus.
VP-14249is similarin lengthandanteroconidshapeto the type Ml shows a shallowalthoughdistinctgrooveon the anteriorside.
ml of B. meadensis.It differs in being wider and in possessing In one of the Ml s the anteroconehas a bilobed shape.Thereis
the anteromedianstylid, a strongeranterolophid,and a stronger a lingual anteriorcingulumin the Mls and M2s, delimitinga
labial cingulum between the protoconidand hypoconid. It is well-developedprotoflexus.All Mls show a strongparastyleand
smaller than the referredspecimensof B. eliasi from Rexroad threeout of five a mesostyle.The M2s have a distinctmesostyle.
Loc. 3 (Hibbard,1956) and lacks a broadanteroconid.The an- Two of the M2s displaya low and thin mesoloph.Lowermolars
teroconeof VP-14247 is less bilobedthanthe Ml's of B. mead- have a strong anteriorcingulum.The mls have a stylid on the
ensis from the Sand Draw and Sandersl.f.s (Hibbard,1972) and latterstructure.Upperand lower thirdmolarsareonly tentatively
much less stronglybilobed than MI's assigned to B. arizonae assignedto this species.
The Onychomyssamplefrom Deer ParkB shows a set of mor(Czaplewski,1987a;Tomida,1987). The m2 differsfrom thatof
the holotypeof B. meadensisin possessinga labialcingulumthat phologicalcharacteristics
thatis not commonin Onychomysfrom
extendsaroundthe posteriormarginof the hypoconidand in not otherlocalities.Amongthem is the presenceof styles andstylids
havingthe posteriorcingulumexpandedmediallyto form a pos- that, althoughpresentin othersamples,are not as frequentas in
teroconulid.A posteroconulidis presenton specimensof B. ari- the molarsfromDeer ParkB. The taxonomicvalueof theseextra
zonae fromthe 111 Ranchl.f. (Tomida,1987). It is similarto the cusps is not known and we do not feel confidentnow to name a
specimenof B. arizonaefrom the VerdeFormationillustratedby new species on the basis of this occurrence.
Carletonand Eshelman(1979) revised the fossil Onychomys
Czaplewski(1987a, fig. le).
When Hibbard(1956) describedthe faunaof the 'MeadeFor- species, includingthose from the Meade Basin. However,they
mation' (=Ballard Formation),no sigmodontineswere reported based theirconclusionsto a greatextenton mandibleswith comfromDeer ParkA. Fromthe overlyingSanders1.f.at Big Springs plete tooth rows. Measurementsand descriptionsof the upper
Ranch,Hibbard(1956) describeda new species of Bensonomys, molarswerenot included.Comparingthe materialfromDeerPark
B. meadensis,the only species of Bensonomysthathe listed from B with those measurementsin commonobtainedby Carletonand
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MARTINET AL.-BLANCAN RODENTS AND LAGOMORPHSFROM KANSAS
1083
TABLE8-Measurements of Onychomyscf. hollisteri molars from Deer Park
B. Abbreviationsas in Table 4.
N
3
12
ml
m2
m3
Ml
M2
M3
ISM
4
5
2
6
3
5
3
1
Length
Min. Mean Max.
1.82
1.33
0.98
1.88
1.36
1.42
1.03
1.96
1.39
0.79
1.90
1.50
1.07
2.07
1.44
Width
s.d.
N
Min. Mean Max.
0.06
0.05
0.07
0.04
5
6
2
5
3
1
0.96
1.00
0.86
1.11
1.10
-
1.12
1.20
1.13
0.84
1.12
1.22
0.89
1.31
1.18
s.d.
0.09
0.07
0.04
6
leading to the modem 0. torridus, until we confirm this hypothesis with further study, we will temporarily maintain 0. hollisteri
as a distinct species rather than as a chronomorph of 0. torridus.
8
10
9
11
12
4
15
13
FIGURE
9-Onychomys cf. hollisterifromDeer ParkB. 1, R Ml (FHSM
VP-14212);2, R Ml (FHSMVP-14431);3, R Ml (FHSMVP-14432);
4, R Ml (FHSMVP-14433);5, R M2 (FHSMVP-14215);6, R M2
(FHSMVP-14411);7, R M3 (FHSMVP-14367);8, L ml (FHSMVP14437); 9, R ml (FHSM VP-14209); 10, L m2 (FHSMVP-14213);
11, L m2 (FHSMVP-14434);12, R m2 (FHSMVP-14435);13, L m2
(FHSMVP-14436); 14, L m3 (FHSMVP-14365);15, L m2 (FHSM
VP-14366).Scale bar = 1.0 mm.
Eshelman (1979), shows that there are similarities between the
Deer Park B Onychomys and both Onychomys hollisteri from the
Borchers 1.f. and Onychomys gidleyi from a number of localities.
The mls from Deer Park B are the same length as those of O.
gidleyi, but they are relatively narrower (Table 8). This difference
is also present in the m2s. The m3s of 0. gidleyi are larger than
the one m3 from Deer Park B that is tentatively assigned to Onychomys.
Onychomys hollisteri from Borchers has an m2 of similar size
to those from Deer Park B, but the Deer Park B mls and m3s
are relatively larger. The size of the Deer Park B Onychomys m3s
fit the trend towards smaller m3s in the lineage 0. bensoni-O.
hollisteri-O. torridus proposed by Carleton and Eshelman (1979),
being intermediate between 0. bensoni from the Benson 1.f. and
0. hollisteri from the Borchers 1.f.
Onychomys material from the White Rock 1.f., described by
Eshelman (1975) as Onychomys. sp., is small and was later assigned to 0. hollisteri by Carleton and Eshelman (1979). Its measurements are similar to the material from Deer Park B (Table 8).
In addition to the metric similarities, both samples have Ml with
an anterior infolding, which is a character not usually present in
other Onychomys species (Eshelman, 1975).
Although, as noted above, Carleton and Eshelman (1979) recognized 0. hollisteri as an intermediate in the phyletic sequence
SIGMODON
MINORGidley, 1922
Figure 10
Material examined.-FHSM VP-14171, nine mis; 14172, three
m2s; 14173, four m3s; 14174, two Mls; 14175, three M2s;
14176, three M3s; 14177, R ml; 14178, two R mls; 14179, six
m2s; 14180, four m3s; 14181, seven Mls; 14182, two M2s;
14183, three M3s; 14420 R ml; 14421-2 two LMls; 14423 R
M1.
Discussion.-The genus Sigmodon has a long history in the
Meade Basin, beginning in the early Blancan with Fallen Angel
B (Martin et al., 2000). Cantwell (1969) and Martin (1979, 1986)
synonymized S. medius with S. minor, and in the Meade Basin
we currently recognize two chronomorphs, S. m. /medius for some
Blancan localities prior to Borchers, and S. m. Iminor for the
Borchers population. Molars of S. m. Iminor tend to be shorter
and narrower than those of S. m. /medius (Martin, 1979), and this
can be readily seen in the illustrations (Fig. 10). Lower molar
measurements of the Deer Park B S. minor generally fall within
the range of those published earlier for S. minor Imedius (Martin,
1979; table 1, as S. medius) with the exception of m3 length,
which is more similar to the same measure in S. m. /minor (Table
9). We do not know yet if this difference represents the beginning
of a dwarfing trend during the late Blancan or is a population
feature peculiar to the Deer Park B sample.
1
67
5
FIGURE
views
of
and
lower
firstmolarsof late Blanupper
10--Occlusal
can Sigmodonfromthe MeadeBasin.S. minorImedius(DeerParkB):
1, FHSMVP-14420,R ml; 5, FHSMVP-14421,L M1; 6, FHSMVP14422, L M1; 7, FHSMVP-14423,R Ml. S. minor/medius(Rexroad
Loc. 3); 3, FHSM VP-14438, L Ml. S. minor /minor (Borchers); 2,
FHSM VP-14439, R ml; 4, FHSM VP-14440, L Ml. Scale bar = 1.0
mm.
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1084
JOURNALOF PALEONTOLOGY,
V. 76, NO. 6, 2002
TABLE
of SigmodonminormolarsfromDeerParkB. Ab9--Measurements
breviations
as in Table4.
Measurements similar to those made on the M2 were taken
from the remaining teeth (Table 10). Measurement data from Deer
Park B were compared with those from Rexroad 3, the type loWidth
Length
cality of Neotoma quadriplicata, by means of t-tests at the 0.05
N Min. Mean Max. s.d.
N Min. Mean Max. s.d.
level of significance. Three significant differences were found; the
ml 7 1.91 1.99 2.08 0.07 13 1.23 1.38 1.56 0.10
greatest occlusal width of Ml and the distance between the base
m2 7 1.31 1.51 1.61 0.11
7 1.45 1.55 1.61 0.06
of the crown and the base of the mesolingual fold on both ml
m3 7 1.61 1.79 1.95 0.15
7 1.33 1.48 1.56 0.08
and m2. In the former case, the population from Deer Park B is
1.54 1.67 1.85 0.12
M1 8 1.96 2.16 2.33 0.14
wider; in the latter case, the fold is deeper in the Deer Park samM2 4 1.40 1.44 1.50 0.05
4 1.38 1.52 1.64 0.13
4 1.23 1.45 1.61 0.16
M3 4 1.39 1.48 1.57 0.07
ple. Zakrzewski (1991, 1993) showed that although folds tend to
be deeper in more advanced woodrats, they are also highly variable characters in Blancan populations. For example, all the folds
on ml of N. leucopetrica from the White Rock l.f. are signifiNEOTOMA
QUADRIPLICATA
(Hibbard, 1941b)
cantly deeper than those of N. quadriplicata from Rexroad Loc.
Material examined.-FHSM VP-14253-14260, eight mls;
3; however, there are no significant differences in fold depth on
14261-14265, five m2s; 14266-14275, 10 m3s; 14276-14279, the m2s and the anterolingual fold on m3 is significantly deeper
four Mis; 14280, M2; 14281, M3; 14282, 29 partialmolars.
in N. quadriplicata. Likewise, N. leucopetrica was significantly
Discussion.-WoodratteethfromDeer ParkB have the typical different from N.
quadriplicata only in length and width of ml
occlusalpatternfoundin otherBlancanpopulations.This includes and m2. It may be of interest that no significant differences were
the staggered mesolophid on ml and m2 (Zakrzewski, 1993), and found
previously in Ml among Blancan woodrats from Kansas
the S-shaped m3. Two mls (VP-14255, 14256) have an antero(Zakrzewski, 1991, table 2). Whether the differences seen belingual fold and two (VP-14253, 14257) have an enamel islet on tween the Deer Park B and Rexroad Loc. 3 samples are a function
the anterolophid. Three m3s (VP-14267, 14268, 14271) have a of
sampling or reflect an evolutionary change will require addistylid at the posterior external edge of the mesolophid. All Mis
tional study.
have a shallow anterolingual fold and three major roots with a
centrally located accessory root. VP-14277 has additional accesFamily ARVICOLIDAE
Gray, 1821
sory roots between the central accessory and posterobuccal root.
NEBRASKOMYSREXROADENSISHibbard, 1970
The M2 (VP-14280) represents an immature individual. MeasureFigure 11
ments (in mm) of the M2 are greatest length = 3.02 mm, greatest
Material examined.-FHSM VP-14076, R ml; 14082, R m2;
width = 1.91 mm, distance between base of crown (bc) and an14087, L m3; 14091, L MI; 14098, two L M3s; 14108, L ml;
terolingual fold = 1.51 mm, bc and anterobuccal fold = 0.54
mm, bc and mesolingual fold = 0.78 mm, bc and posterobuccal 14111, L ml; 14112, L ml; 14131, L M3; 14132, three molar
fold = 0.54 mm. Accurate measurements could not be obtained pieces; 14133, R M3; 14134, two M3 pieces.
from the M3 because of its eroded condition.
Discussion.-Nebraskomys is a small arvicolid (Table 11) with
TABLE10--Measurements (in mm) of Neotoma quadriplicatamolars from Deer Park B and Rexroad Loc. 3, respectively. N = number of specimens, i =
mean, s = standarddeviation, OR = observed range, GL = greatest occlusal length, GW = greatest occlusal width; remaining abbreviationsdenote
distances between base of crown and base of folds: ab = anterobuccal,al = anterolingual,ml = mesolingual, pb = posterobuccal,pl = posterolingual.*
= significantdifference at p < 0.05.
Rexroad Loc. 3
Deer Park B
N
ml
GL
GW
pl
pb
ml*
ab
m2
GL
GW
pl
pb
ml*
ab
m3
GL
GW
pb
ml
s
OR
N
s
OR
6
5
6
5
6
5
3.4
1.85
0.96
0.83
0.79
0.91
0.16
0.3
0.16
0.17
0.07
0.13
3.15-3.66
1.43-2.13
0.74-1.16
0.68-1.11
0.71-0.92
0.65-1.27
30
30
30
30
30
30
3.39
1.86
1.09
0.88
0.92
0.94
0.18
0.15
0.18
0.09
0.12
0.18
2.96-3.80
1.56-2.13
0.80-1.38
0.76-1.12
0.67-1.16
0.62-1.27
5
5
4
5
5
4
3.1
2.11
1.15
0.73
0.7
1.12
0.11
0.51
0.23
0.16
0.09
0.09
2.94-3.21
1.24-2.57
0.85-1.35
0.48-0.90
0.62-0.82
1.01-1.23
10
10
10
10
10
10
3.06
1.88
1.31
0.87
0.9
0.98
0.18
0.13
0.15
0.07
0.77
0.15
2.78-3.29
1.68-2.13
1.10-1.62
0.76-0.96
0.77-1.05
0.71-1.29
8
9
8
8
2.48
1.91
0.78
0.79
0.26
0.25
0.14
0.23
2.12-2.47
1.47-2.14
0.62-1.04
0.51-1.29
10
10
10
10
2.39
1.91
0.68
0.71
0.13
0.14
0.11
0.15
2.24-2.63
1.62-2.09
0.49-0.85
0.34-0.85
M1
GL
GW*
4
4
3.72
2.5
0.14
0.25
3.51-3.83
2.28-2.81
10
10
3.58
1.96
0.26
0.18
2.96-3.86
1.68-2.34
al
ab
4
3
0.92
0.68
0.19
0.11
0.78-1.19
0.57-0.79
10
10
0.98
0.7
0.18
0.07
0.73-1.25
0.57-0.84
3
0.78
0.11
0.67-0.89
10
0.74
0.1
0.62-0.89
4
0.74
0.11
0.64-0.87
10
0.7
0.07
0.57-0.84
pb
ml
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MARTINET AL.-BLANCANRODENTSAND LAGOMORPHS
FROMKANSAS
TABLE11-Measurements of Nebraskomysrexroadensis molars from Deer
Park B. Abbreviationsas in Table 4.
K
4i
3
~
1 ~2
1
7
~19
5
1085
6
m
FIGURE 11--Lower firstmolarsandupperthirdmolarsof Nebraskomys
rexroadensis
fromDeerParkB. 1, 2, FHSMVP-14108,Lml; 3, 4,
FHSMVP-14112,L ml; 5, 6, FHSMVP-14111,L ml; 7, 8, FHSM
VP-14131,L M3;9, 10,FHSMVP-14133,RM3.Scalebar= 1.0mm.
a very simple dentitionand thick, undifferentiated
enamel.Originally describedby Hibbard(1957) from the Sand Draw 1.f.,the
genus includes two species, N. mcgrewi and N. rexroadensis (Hib-
bard, 1970). Accordingto Hibbard(1970), N. rexroadensishas
shorterdentine tracts than N. mcgrewi and T1-2 on m2 in N.
mcgrewi are more directly opposite than in N. rexroadensis. A
quantitativecomparisonof dentinetractheightin the two species
was not made for this project,but the overallmorphologyof the
Deer ParkB specimensbest matchesthatof N. rexroadensis.
Althoughsamplinghas been ratherintense for Blancantime,
Nebraskomyshas prevoiouslybeen recordedin the MeadeBasin
only at RexroadLoc. 3 (Hibbard,1970). It is also present at
Bender ic (UMMP 60930, Rm2; new record)and now at Deer
ParkB. Its absencefrom Sanders(Hibbard,1956), within 10 km
from the Deer Parksite, is consideredsignificant,and this negative evidence,in concertwith otherrecords(Table13), helps solidify the idea that Sandersis youngerthanDeer Park.
As suggestedby Patton(1965) and Hibbard(1970), and confirmedby Winklerand Grady (1990), Nebraskomysappearsto
have been ancestralto the middle PleistoceneAtopomysof the
eastern United States. The origin of Nebraskomysremainsobscure,thoughits simple schmelzmuster(von Koenigswald,1980)
and underivedml suggest that it evolved directlyfrom a Promimomysgradearvicolid.
OGMODONTOMYS
POAPHAGUS
Hibbard, 1941
ml
m2
m3
M1
M3
N
Length
Min. Mean Max.
4
1
2
1
3
2.07
1.46
1.23
1.93
1.53
2.16
1.46
1.27
1.93
1.81
2.23
1.46
1.31
1.93
2.00
Width
s.d.
N
Min. Mean Max.
s.d.
0.07
4
1
2
1
3
1.09
1.00
0.84
1.26
1.05
0.07
0.06
0.25
1.19
1.00
0.90
1.26
1.17
1.23
1.00
0.95
1.26
1.23
0.08
0.10
be measuredaccuratelyon two mis from Deer Park A, and is
0.73 and 0.84 mm.
Six of eight (75 percent)Deer ParkM3s have threeroots, the
others have two roots. Thirtyfour M3s from the WendellFox
Pasturelocality displayedthreerootson M3 (Zakrzewski,1967).
Preliminaryexaminationof roots on Ogmodontomys
poaphagus
M3s from a numberof new Blancan sites in the Meade Basin
shows that root numberon M3 is variable,but we do not have a
quantitativecomparisoncompletedat this time.AlthoughHibbard
neverprovideda confirmedstratigraphic
positionfor the Wendell
Fox Pasturesite, preliminarystudyof specimensfromthislocality
in the Universityof Michigancollectionsindicatesit is of middle
Blancanage (Martinet al., 2000). The absenceof Nebraskomys
from WendellFox Pasturesuggestsit was depositedearlierthan
RexroadLoc. 3.
The highestdentinetractson any Ogmodontomys
werereported
by Zakrzewski(1967) froma singleml recoveredfromsediments
in HartDraw on the Big SpringsRanch.Small mammalfossils
referredto the Sandersl.f. were also recoveredhigher in this
section,withinthe type BallardFormation(Hibbard,1956;Bayne,
ml mor1976). Based primarilyon the advancedOgmodontomys
phology, Martinet al. (2000) includedHartDraw at the Ballard
Formationhorizon, but in light of the new stratigraphicinformationabove regardingRexroadLoc. 3, this allocationmust be
consideredonly tentative.
is consideredhereas a distinctNorthAmerican
Ogmodontomys
genus separatefrom Mimomys.No North Americanearly arvicolids with rootedmolarsdisplaynegativeenameldifferentiation
(a synapomorphy for Mimomys), and Ogmodontomys lacks the
well-developedlamellarenamelon triangleleadingedges present
in all Mimomys(von Koenigswald,1980; Martin,in press). Cosomys apparentlysecondarilydeveloped lamellarenamel edges
(von Koenigswald,1980).As the ancestraltaxafor most arvicolid
clades have an ml with a Mimomys-kante,
this feature,as a symplesiomorphy,cannotbe used effectively in the classificationof
early arvicolids(contraRepenninget al., 1990 and Mou, 1997).
Coues, 1876
Family ZAPODIDAE
ZAPUScf. SANDERSI
Hibbard, 1956
Material examined.-FHSM VP-14192 L Ml; 14193 R M1;
14194 L m3.
Figure 12
Discussion.-The latest investigationsof late Pliocene and
Material examined.-FHSM VP-14135, seven molar pieces;
14136, two mls; 14137, two m2s; 14138, two Mls; 14139, four
M2s; 14140, four M3s; 14141, eight mls; 14142, three m2s; TABLE
12-Measurementsof Ogmodontomys
poaphagusmolarsfromDeer
ParkB. Abbreviations
as in Table4.
14143, six m3s; 14144, two Mls; 14145, M2; 14146, six M3s;
14147, nine molarpieces; 14148, R ml; 14402 L ml; 14403 R
Width
M3; 14404 L M3.
Length
N Min. Mean Max. s.d.
N Min. Mean Max. s.d.
Discussion.-In size and morphologythe Deer ParkA and B
0. poaphagusare typicalfor middleand late Blancansamplesof
2 3.08 3.29 3.49 0.29
2 1.85 1.88 1.90 0.04
this species (Table12, Fig. 12; Zakrzewski,1967). Dentinetracts ml
3 1.85 1.98 2.20 0.19
m2
3 1.38 1.51 1.70 0.17
on the labial side of ml (e.g., the hyposinuid)are developedon m3 5 1.60 1.72 1.81 0.08 5 1.18 1.29 1.51 0.14
1
2.66
2.66
2.66
1 1.95 1.95 1.95
boththe Deer ParkA andB specimensto approximately
the same M1
extent as in materialfrom other middle and late Blancansites, M2 5 1.72 2.19 2.45 0.28 5 1.64 1.77 1.91 0.11
such as WendellFox Pasture(Fig. 12). Hyposinuidheightcould M3 6 2.04 2.25 2.46 0.18 8 1.23 1.41 1.54 0.10
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1086
JOURNALOF PALEONTOLOGY,
V. 76, NO. 6, 2002
TABLE13--Comparison of lagomorphsand rodents from some Blancan Meade Basin localities. The modern lagomorphsand rodents of Meade County are
also included. Rex 3 = Rexroad Loc. 3; Ben = Bender lb, Ic (records from Bender lb are new, from our fieldwork. Bender Ic records were obtained
from UMMP collections); DPA = Deer Park A; DPB = Deer Park B; Rex 2a = Rexroad Loc. 2a (including only materialcollected in July, 2001); San
= Sanders. cf = compares favorably.
Ben
Lagomorpha
Leporidae
Rex 3
Hypolagus regalis
Hypolagus voorhiesi
Notolagus lepusculus
Nekrolagus progressus
Pratilepus kansasensis
Alilepus wilsoni
Alilepus sp.
Sylvilagus floridanus
Lepus californicus
Lepus sp.
x
Spermophilus rexroadensis
Spermophilus howelli
Spermophilus cragini
Spermophilus meadensis
Spermophilus tridecemlineatus
Spermophilus spilosoma
Cynomys ludovicianus
Spermophilus sp.
x
x
Dipoides rexroadensis
Procastoroides sweeti
x
x
DPA
DPB
Rex 2a
San
Bor
Mod
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Rodentia
Sciuridae
cf
x
x
x
x
x
x
x
x
x
Castoridae
Geomyidae
Geomys jacobi
Geomys minor
Geomys quinni
Geomyssp.
x
x
x
cf
cf
cf
cf
cf
cf
cf
cf
x
xb, c
x
x
Geomys bursarius
Heteromyidae
Prodipodomys idahoensis
Prodipodomys tiheni
Prodipodomys sp.
Dipodomys hibbardi
Dipodomys ordii
Perognathus gidleyi
Perognathus pearlettensis
Perognathus sp.
Perognathus flavus
Perognathus flavescens
Chaetodipus hispidus
cf
x
x
x
Baiomys rexroadi
Peromyscus kansasensis
Peromyscus hagermanensis
Peromyscus maniculatus
Peromyscus leucopus
Peromyscus sp.
Bensonomys eliasi
Bensonomys meadensis
Symmetrodontomys simplicidens
Onychomys hollisteri
Onychomys gidleyi
Onychomys pedroensis
Onychomys lecucogaster
Reithrodontomys pratincola
Reithrodontomys megalotis
Reithrodontomys montanus
Sigmodon minor
Sigmodon hispidus
Neotoma quadriplicata
Neotoma taylori
Neotoma micropus
x
x
x
x
x
x
x
cf
x
cf
x
x
x
x
x
x
Cricetidae
x
cf
x
x
x
x
cfc
x
x
x
cf
x
cfb
cf
x
cf
cf
x
x
x
x
x
x
x
xb
x
x
x
x
x
x
x
x
x
x
x
x
Arvicolidae
Nebraskomys rexroadensis
Ogmodontomys poaphagus
Ophiomys meadensis
Ondatra zibethicus /meadensis
Ondatra zibethicus /idahoensis
Pliolemmus antiquus
Mictomys landesi
Synaptomys cooperi
Microtus ochrogaster
Zapodidae
Zapus sandersi
Zapus burti
Zapus hudsonius
xc
xb, c
x
x
x
x
x
x
x
x
x
x
x
xc
x
x
x
x
x
x
x
x
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MARTINET AL.-BLANCAN RODENTS AND LAGOMORPHSFROM KANSAS
474
12
112
114 1
1416
13
12-The dentitionof Ogmodontomys
FIGURE
poaphagusfromsome late
Blancanlocalitiesin the MeadeBasin.Deer ParkA: 1) and 2) UMMP
31952(1);from bulk sample,numberedmolarsin separategelatincapsules) (juvenile),L ml; 3) and 4) UMMP31952(2), L ml. Deer Park
B: 5) and 6) FHSM VP-14402, L ml; 15) FHSM VP-14403, R M3;
16) FHSMVP-14404L M3. WendellFox Pasture:9) and 10) UMMP
41432(16), L ml; 11) and 12) UMMP41432(21), L ml; 13) and 14)
UMMP41432(8), R ml. RexroadLoc. 3: 7) and 8) FHSMVP-14441,
L ml. Linea sinuosashownfrom labial side. Scale bar = 1.0 mm.
Pleistocene North American zapodid evolution were provided by
Klingener (1963) and Martin (1989, 1993). Zapodids have been
a constant, albeit rare, constituent of Meade Basin local faunas,
and absence from any fauna is almost certainly due to sampling
bias. Klingener (1963) identified a phyletic relationship extending
from the middle Blancan Z. sandersi of the Meade Basin Wendell
Fox Pasture 1.f. to the modern Z. hudsonius, the latter first appearing in the Mt. Scott l.f. (Rancholabrean), also from Meade
County. The Borchers Zapus sample, accepted by Klingener
(1963) as the somewhat aberrantZ. burti, presents a troublesome
break in an otherwise simple lineage, and suggests the taxonomy
of Meade Basin Zapus should be reviewed.
Length of the m3 = 0.87 mm; width = 0.77 mm. Length of
the two Mis is 1.23, 1.29 mm and width is 1.14 and 1.18 mm.
DISCUSSION
Age of the Deer Park Assemblages.-As discussed above,
among the specimens described from Deer Park A by Hibbard
(1956) was a new species of prairie dog, Cynomys meadensis. In
his monograph on the fossil prairie dogs (and above), Goodwin
(1995) concluded that the Deer Park Cynomys material was relatively modem and intrusive, a position he noted (Goodwin, 1995;
p. 20) that Hibbard had also accepted. Additionally, Zakrzewski
(1967) was concerned because two Ogmodontomys mls from
1087
Deer Park A appeared to lack well-developed dentine tracts as
found on specimens from the presumably older Bender 1.f. Zakrzewski mentioned the possibility of reworking older material
by the artesian system at Deer Park A. We re-examined the Ogmodontomys specimens from Deer Park A and, as noted above,
the dentine tract heights are within the range of those from Wendell Fox Pasture (Fig. 12), the latter which is likely older than
both Deer Park and Bender. We did not recover any specimens
from Deer Park B that suggest reworking or mixing with younger
units. Additionally, while it seems clear there is a time differential
between Deer Park A and B simply based on superposition, this
difference is probably not pronounced, as the species common to
both levels show no obvious morphological disparity.
There is a growing body of evidence to confirm Hibbard's
(1972) placement of the Deer Park depositional regime in the
middle or late Blancan [for many years Hibbard had tried to sequence some of the southwestern Kansas local faunas we now
know to be Pliocene (e.g., Dixon, Deer Park, Sanders; Hibbard,
1956) into a Pleistocene glacial-interglacial sequence, but by the
time he and Morris Skinner published their monograph on the
Sand Draw 1.f., Hibbard came to realize many of these faunas
preceded the first evidence of continental glaciation]. According
to Lindsay et al. (1975) the sediments at Bender (presumably Loc.
Ic), Rexroad Loc. 3, and Sanders are normally magnetized. All
were likely deposited during the Gauss chron, between about 2.63.6 Ma. It is not possible now to determine their temporal placement with much finer resolution, although in Figure I we provide
a tentative chronological hypothesis. These localities appearto be
younger than a series of Blancan localities including Keefe Canyon, Raptor IC, Ripley, and Fox Canyon (Martin et al., 2000).
The sediments at Fox Canyon are magnetically reversed (Lindsay
et al., 1975) and have been assigned to the Gilbert Chron, perhaps
about 4.3 Ma (Martin et al., 2000).
Paleoecology.-Hibbard (1956) reported the presence of flour
sand tubes in his quarries at Deer Park A. He noted that many of
the fossils appeared to have been churned, and larger mammalian
teeth often acted as "millstones." Although this may well have
been the case, much of the small mammal material from both
Deer Park A and B is in pristine condition, showing no signs of
tumbling or significant transport. The material from locality A is
mostly white, confirming the dominant presence of sand at the
locality. Fossils from level B range in color from white to dark
reddish brown (Munsell soil color chart). What damage is observable in rodent teeth from both localities seems to have been
primarily produced by carnivore gastric acid digestion (Andrews,
1990). There is some additional damage that may have been
caused by plant roots and humic acids (etched traces and white,
sloughed edges). In particular, the material from level B shows
more modification in this regard, as the sediments from B are
within the modern soil zone.
We agree with Hibbard (1956) that the Deer Park A material
was deposited in an active spring. The absence of Ondatra zibethicus /meadensis, Procastoroides sweeti, Pliolemmus antiquus,
and large mammals in general from level B is probably not accidental. In particular,beavers and muskrats, always rare in Blancan localities of the Meade Basin, were likely restricted to environments with sufficient water to satisfy their foraging, reproductive, and nesting strategies. Perhaps Pliolemmus preferred very
mesic environments, or was at least semi-aquatic. By Deer Park
B time, the energetic spring at that location was probably gone,
replaced by a limited swampy region with minimal standing water. The fine sediments of Deer Park B do not suggest the presence
of a stream with any competency, if one was there at all. A spring
exists now in Meade State Park less than 0.25 km north of the
Deer Park quarries. This spring could represent the northward
continuation of an artesian system that may have been represented
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1088
JOURNAL OF PALEONTOLOGY,V. 76, NO. 6, 2002
in the Rexroad Formation (earlier in Blancan time) by the Rexroad 2 and 2a localities, both of which occur in the same pasture
about 0.25 km south of Deer Park (Stevens, 1965).
The small mammals from Deer Park B in aggregate suggest an
oldfield or prairie ecosystem that might have been somewhat more
arid than that in the area today. Forest dwelling animals (such as
the tree squirrels) are absent. There is not a large enough fossil
sample to make any meaningful statistical comparisons among the
species represented, but the relatively large number of pocket gophers of the genus Geomys certainly attests to the proximity of
relatively deep, loose soil or sand (Davis, 1966; Schwartz and
Schwartz, 1981). The presence of Baiomys (not currently found
in Kansas) and the absence of Reithrodontomys is interesting,
though it is difficult to know if this replacement has any ecological significance in itself. Based on preliminary study only,
Baiomys appears to be more common than Reithrodontomys
throughout the Blancan in the Meade Basin. Local faunas with
exclusive representation of Reithrodontomys, as in the modem
biota, are seen only from Borchers time (2.1 Ma) onward. Both
genera are restricted to open, grassy environments. Baiomys is
limited in its distribution today to the southwestern United States
and Mexico. According to Davis (1966), in Texas Onychomys
torridus prefers open sandy, gravelly topography with sparse vegetation. If the Deer Park B grasshopper mouse is correctly interpreted as a member of the lineage leading to 0. torridus, then
this occurrence, with a species of Baiomys, suggests a more arid
ecosystem than now present in Kansas.
Not counting aquatic taxa, the number of rodent species from
Deer Park B (18) is almost the same as that in southwestern Kansas today (17), lending more evidence to the conclusion reached
earlier by Martin and Fairbanks (1999) that diversity, represented
by species richness, remains fairly stable in Meade Basin rodent
local faunas, despite considerable species turnover. Two species
in the Deer Park B 1.f. and three species in the modem fauna are
arvicolids. Although there is considerable variation in the data,
Montuire et al. (1997) demonstrated a significant relationship between a number of temperature parameters and number of arvicolid species in modern global faunas. Based on these data, the
mean annual temperature during Deer Park B time could have
ranged between about 9 to 21 degrees Celsius; within current
values. Differences in frequency of other taxonomic groups between the modem rodent fauna and that of Deer Park B may at
least partly represent sampling or taxonomic bias. For instance,
there are two species of Perognathus in southwestern Kansas today, whereas only one species is recognized from Deer Park B.
It may be that the fossil sample from Deer Park B was not large
enough to represent a second species of pocket mouse, or molars
are so similar in small pocket mice that with the limited sample
available a second species cannot be recognized. In any case, the
combined rodent species from Deer Park B suggest a prairie ecosystem that might have been slightly more arid than that found
today in southwestern Kansas.
ACKNOWLEDGMENTS
We are most appreciative of the continued help and good cheer
provided by the staff at Meade State Park, including M. Goldsberry, J. Lillie, C. Helms, and J. Zuercher. G. Gunnell and P
Gingerich provided loans of many important University of Michigan specimens. This research was supported by grants 5963-97
and 6547-99 from the National Geographic Society and matching
funds from MSU.
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