• TABLE OF CONTENTS
HIDE
 Title Page
 Synopsis
 Introduction
 Acknowledgement
 Abbreviations
 Stratigraphy
 Distinguishing characteristics...
 Systematic descriptions
 Age and correlation
 Summary and conclusions
 Literature cited
 Back Matter






Group Title: Bulletin of the Florida State Museum
Title: Miocene and Pliocene artiodactyls, Texas Gulf Coastal Plain
CITATION THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00001524/00001
 Material Information
Title: Miocene and Pliocene artiodactyls, Texas Gulf Coastal Plain
Series Title: Bulletin of the Florida State Museum
Physical Description: 116-226 p. : illus., map. ; 23 cm.
Language: English
Creator: Patton, Thomas Hudson, 1934-
Publisher: University of Florida
Place of Publication: Gainesville
Publication Date: 1969
 Subjects
Subject: Artiodactyla, Fossil   ( lcsh )
Paleontology -- Pliocene   ( lcsh )
Paleontology -- Miocene   ( lcsh )
Paleontology -- Texas   ( lcsh )
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Bibliography: p. 220-226.
General Note: Cover title.
 Record Information
Bibliographic ID: UF00001524
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0834
notis - ACK3572
alephbibnum - 000442921
oclc - 00134116
lccn - 71628702

Table of Contents
    Title Page
        Title Page 1
        Title Page 2
    Synopsis
        Page 115
    Introduction
        Page 116
    Acknowledgement
        Page 117
    Abbreviations
        Page 118
        Page 119
    Stratigraphy
        Page 120
        Page 121
        Page 122
        Page 123
        Page 124
        Page 125
    Distinguishing characteristics of the faunas
        Page 126
        Page 127
        Page 128
        Page 129
        Page 130
        Page 131
        Page 132
    Systematic descriptions
        Page 133
        Page 134
        Page 135
        Page 136
        Page 137
        Page 138
        Page 139
        Page 140
        Page 141
        Page 142
        Page 143
        Page 144
        Page 145
        Page 146
        Page 147
        Page 148
        Page 149
        Page 150
        Page 151
        Page 152
        Page 153
        Page 154
        Page 155
        Page 156
        Page 157
        Page 158
        Page 159
        Page 160
        Page 161
        Page 162
        Page 163
        Page 164
        Page 165
        Page 166
        Page 167
        Page 168
        Page 169
        Page 170
        Page 171
        Page 172
        Page 173
        Page 174
        Page 175
        Page 176
        Page 177
        Page 178
        Page 179
        Page 180
        Page 181
        Page 182
        Page 183
        Page 184
        Page 185
        Page 186
        Page 187
        Page 188
        Page 189
        Page 190
        Page 191
        Page 192
        Page 193
        Page 194
        Page 195
        Page 196
        Page 197
        Page 198
        Page 199
        Page 200
        Page 201
        Page 202
        Page 203
        Page 204
    Age and correlation
        Page 205
        Page 206
        Page 207
        Page 208
        Page 209
        Page 210
        Page 211
        Page 212
        Page 213
        Page 214
        Page 215
        Page 216
    Summary and conclusions
        Page 217
        Page 218
        Page 219
    Literature cited
        Page 220
        Page 221
        Page 222
        Page 223
        Page 224
        Page 225
        Page 226
    Back Matter
        Page 227
Full Text




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MIOCENE AND PLIOCENE ARIIODACTYLS.
T EXAS GUELII- COS IAL PLAIN


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FI'rcpE 13. Aepyc'nelwu sp. '"FTBEC; : 118:3-46, Type ,;
front and lateral views. B. right r-idiils-ii3na,


A. right netLatarsal,
rear and lateral views


partial dentition. In thei characters of the mandible and lower molars
listed below, Plrotolabiis cicel resembles Proca)mlu.s, but is (istitict
from it iii the configturation o f tie skull, upper incisors, lower pre
molars,,and itmtapodiai.s. In his paper on the Burge and Minnrchi -
duza fatl nas, \\ebb (1.969) revises the generic dlia gnosis of Proto-
lobis as follows:
)erititlifi 1,'iTCUilI.'e.d wVirh flIrItitJuIJ ,,ii-j.x-r n ciszm. n111 13!.__rT itidVI idi._Lti
P. double rb'teed (:heck t'cet-i prtNportiin.rlt llv t.i.rr ower th ; n ii n fPr nmra h-c.:. ;
i;-arir( "v r jrid- higher lcr.i ic rld tha~l Ira Mtwo i.L. P o.'t-ii .t- ,fri .1 Jf P. Ln~rr)iW ii
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199 PATTON. TEXAS ATIODACt~YS 1S1

rotnsu i Deiduous ehek teeh preopotmonsly 'ueh smller and lo'r
rosned iloina Ploeotel.a dp. sad dye evey steedas ond leieg lIngual in-
flcns. Deep casnonis witb Esled baeise udalu sad 0 shaped aygonaoi c ec-
Limbe about bhal long tu In Roaomebu gnd and create gy light; eta-
podlals unused in mate. individuals
In a discussion of the confusion In the synomy of the specie of
Prothbbis, Webb (199) suggests that two species should peently
be recognized: "a mailer form which should probably be called
Proolabi fissides but is best represented by the material Matthew
named P. ongiceps, and a larger form presented by Potolabis
heterodonta and its synonym P angustides" I bevef that other
evidsee indicteas a necessity for continued separaion of P. fsideno
and longceps: Cope's (1874 a: 327) unMllustated description of Pro
amelu fisidens points out t thte reduton of the lower premolars
approximates that of Procumel occidenta= and Matthew (1901)
states that the premolars of P. fiside are the least reduced of the
procameline species In comparison, the prmolars of Prooloabi long-
ceps (=P. montanus) ar quite reduced, P and Pe (although Px
loss is probably sre) being absent. This suggests that although the
genus Protoabis may contain twnosize groups, P, longieps and
P. fisstdens should be considered distinct In addition, Webbs gen-
















PFnSS 14, PtdotiWs iantaoi s n. sp (UTBEG 31133-33M Type); ri
"ramu, tabtl nve of ansus sad P.-L., ocdeal view at eoli fou
P.-. ead P.-...

.1 C. ,








eric diagnosis should be augmented to include mention of this pre-
molar reduction.
The molars of UTBEG 31132-332 from the Lapara Creek Fauna
are 11-25 per cent smaller than those of P. heterodontus from the
Burge and Minnechaduza Faunas. The Texas specimen is about 10
per cent larger than P. longiceps. Because of differences in overall
size, shape, and proportions of the lower premolars, I am separating
the Gulf Coast specimen from the Great Plains forms as a distinct
species, Protolabis notiochorinas. This is the only species of Protolabis
now known from the Gulf Coastal Plain, as I have referred P. francisi
Hay to Oxydoctylus benedenttus (page 135).

Protolabi notiochorinos new species
Figures 14, 15, Table 11
TYPE. UTBEG 31132-332 mandible with P4-Ma intact; P2-Pa
broken below crown. P. alveolus present.
TYPE LocAI rr Bridge Ranch; Site 17, Bee County, Texas
OCCURENCs. Lapara Creek Fauna.
REtERaE MATEaL-UTBEG 31081-261, -293, -612, -525, UTBEG
31132-319, all metapodials.
ETrrorlO notios = Gr. southern; chores = Gr. land, country.
DIAcxOSs. A species of Protolabti intermediate in size between
P. longiceps and P. heterodontus. Premolars more reduced than n
heterodonlus. P, two-rooted; Ps, Pu, and P4 highly reduced. P4 very
narrow and with posterior lobe smaller than middle lobe, m contrast
to longiceps and heterodontus. Masseteric region of mandible more
greatly flared than in longiceps.
DSCRIrrToN. The shape of the mandible of P. notiochorinos
appears to resemble closely P. longiceps from the Pawnee Creek,
except that the masseterie region of the Texas specimen has a greater
posteroinferior flare The mandibular foramen is situated below the
occlusal level, and its inferior border is interrupted by a wide, 4-6-mm
groove extending down and forward for about 28 mm, probably
serving to receive the mandibular artery and nerve. A smaller foramen
appears toward the distal end of this groove on the left ramus. Only
a hint of this foramen is present on the right ramus. A mental fora-
men occurs in the lower third of the ramus jist below the posterior
border of Mi. The ranm are broken anterior to P1. P1 of the left ramus
is only party intact, but shows evidence of an alveolus from an
anterior rootFaus establishing the pessence of a two-rooted P. P,








Table 11 Mua s umwrs orF TYPE or Frookbs not ohoruio, Nzw SpecTm
Lower jaw UTBEG 3112-32
Depth of namua below center of M1 51.5
Depth of amus below center of Mi 40.2
Depth below center of P,-P. diasmea 0,0
Length, P,-P, diatens 26.2
P.M, 112.0'
P.-M, 95.0
P.PI 2.7.0
M-M, 84.01
PK length x width 8.0
P, length X width 113 X 5.7
MP, length X width 172 X 11.1
M. length X width 27.0 X 150
M., length X width 4.0 X 18.0
'Appxleml.a

and Pt are separated by a rather short (26 mm) diastema. Of the
premolars, only the right P. is preserved. The roots of Ps and Pa are
visible and reveal the highly reduced condition of these teeth. P4 is
short anteroposteriorly and extremely narrow (5.7 rm). In com-
parison to Procameiws, the anterolingual flexid of PF is only weakly
expressed. A faint posterolabial fold is visible. The most notable
feature of P4 is that the posterior lobe is narrower than the anterior,
in contrast to other camels, in which the wedge-shaped P4 is much
wider posteriorly than anteriorly. The absence of any elaboration of
the posterior lobe precludes the formation of a posterior fossettid.
The unusually narrow posterior lobe of P. of P. notiochornos recalls
a similar construction of Ps in P. heterodontus. Also in P. heterodontus
P4 is marked by a weakly inflected anterior lobe. The molars of P.
notiochorinos are quite narrow but simple, with only faint expression
of lingual ribs or stylids. The only complete uncoossified camel meta-
podial (UTBEG 31132-319) from the Lapara Creek Fauna is only
194 mm long, but a broken specimen (UTBEG 31081-293) appears
to be approximately 20 per cent larger.
DIscU sSon The Lapara Creek specimen resembles P. hetero-
donfus from the Burge and Minnechaduza Faunas in many respects,
and could well be a close relative of it. The relatively more advanced
stage of evolution represented by P. rnotochorinos as compared to P.
hterrodonfus suggests ,an ancestor-descendet phylogenetic relation-
ship, but the inference is refuted by the presence of these two forms
in contemponeous deposits (Clarendonian). More probably, P. no.
tiochorinos rereents a branch off an early P. heterodontus stock and
,?






























FIcUen 15 Protolbis oioc no, n sp. (UrBEG 118232, Type), man-
dible, labial view of right ramus, occusal .ew of both rami,

one that evolved separately at a more accelerated rate than the Great
Plains species, but the synonymy and systematic relationships of the
various species will remain uncertain until this genus is thoroughly
reviewed.

Procamels Leidy, 1858
Procamelus is among the most abundantly represented artiodactyl
groups recovered from the faunas described in this paper. Although
absent from the Garvin Gully and Burkeville Faunas and only spar-
ingly represented in the Cold Spring, it is the most numerous form
in the Lapara Creek Fauna. Whereas Oxydactylus is the commonest
camel in the Garvin Gully Fauna, Procamelus assumes this position in
the Lapara' Creek Fauna
The type species is Procamelus occidentalis, described by Leidy
(1858) from the late Miocene deposits of the Niobrara River Valley
in Nebraska Procamelus is represented m the Texas faunas by 12
well-preserved mnandibles, 12 metapodmals, 1 humerus, and 3 radius-
ulnae. Many of the nominal species formerly assigned to the genus








Procamelus are now regarded as nomina nuda or nomena dubia
(Gregory, 1942, Webb, 1969). In general, three species groups of
Procamelus are currently recognized (Gregory, 1942; Webb, 1969):
P. occidentali and its probable descendant P. grand; the wide-
toothed, short-jawed P. robustus; and the group of small procamelines
composed of P. graodis, P. leptognathus, and P. coariaus. Procmelus
specimens from the Lapara Creek Fauna are assignable to two distinct
species: P. occidentalis Leidy and P. grandis Gregory. This bipartite
division of Procamebse in the Lapara Creek Fauna is discernible in
the size distribute and moology of the and mopholoof the mandibles and the meta-
podials.












2 cm

FICUiz 16 Procmef occdenals (TBEG 81081-065); left ramu, labial
view Ii-M,. occlusal view P.-M..


Procamels occidental Leidy, 1858
Figure 16, Table 12
TwrP. USNM 797, right Pa-Ms (broken), fig. Leidy, 1869, PV,
fig. 5.
OCcuRRENCE Lapara Creek Fauna.
RE E MATERAI. Mandibles: UTBEG 31081-665, 31081-738,
31081-49, 31081-313, 31081-241, 30896-475, 31170-6; humerus: UTBEG
31081509; radus-ul UTBEG 31081-582, -451, 31132-18; meta-
carpals: UTBEG 31081-418, -503, -840,'-1080, -1085, metatarsals:
UTBEG 31081-407, -408, -417, -438, -490, -159
D.SCRi ON. The smaller of the two Procanelus species from
the Lapara Creek Fauna, P. occidental is the best represented of








all the camels in numbers of specimens. UTBEG 31081-665 (Fig. 4B)
consists of both rami of the mandible with one ascending ramus
virtually complete and the dental series intact, except for the incisors,
which are present but broken on the left side and missing on the
right. The lower jaw and lower dental series show few or no quali-
tative differences between the Lapara Creek specimens and either
P. occdentalis or P grandis The lower incisors are too fragmentary
for detailed description. The ventral surface of the mandible shows
no deep sulcus along the symphysial plane between left 11 and right I:,
as occurs in Pliancheni and Nothotylopus A short diastema (10 mm
between alveolar borders) is present between Is and /C. The anterior
surface of /C is marked by a very thin, bladelike flange which is
pinched off sharply from the body of the tooth. A similar flange is
present, but much less developed in Pt Less defined flanges occur
along the posterior edges of /C and Pt. The prominent single-rooted
P, lies roughly midway between /C and P2. The Pt-P2 diastema, as
mentioned, Is almost equal to that between /C and Pi, a condition
characteristic also of Protolabis, but in Nothotylopus the PI-Pa di-
astema is more than twice the distance between /C and Pi.P2 of P.
occdentalis is thin, sharp, and considerably reduced in comparison to
Ps and P4. The two posterior premolars resemble one another closely

Table 12 Mr.Asu'Mnrs oF Procamefus occidenta a
UTBEG UTBEG UTBEG
Lower jaw 31081-665 8089-475 31081-241
Condyle to angular process 20.4
Length of /C-P diastema 228
Length of PrP, diastema 80,1
Depth of ramus below
center of MK 43.2 39.1 435
D .pth below center of M 85.2 80.0
Depth below center of M2 30
Deph below center a
P-P. diastrnm 27.6 242
PVM, 1157 112,0' 1184
Pr-M 108.0 102.0 100.5
P.FP 83.6 34.8U 37.0'
Pr-P 26.2 262' 28.4
Mi-M 88.1 77.0 810
P', length X wdth 94 X 5.0
P., length x width 12.7 x 60 18.5 x 5.8
P,, length X width 14.0 x 8.5 15.0 x 7.4 159 x 8.0
M, length x width 20.1 x 12.8 20.- X 12.0 210 x 12 5
Mi length X wV& 260 X 15.0 237 X 123 25.6 X 137
Ma. length xw 38.1 X 15.0 3486 13.8 80.6 X 14.2
sn








in cusp morphology. They are marked by a pronounced anterolingual
fold (lexid) between the anterior (paraconid?) and middle meta-
stylid?) lobes. The lingually flexed paraconid makes a slight dogleg
anteriorly, a feature also observable in some species of Pronioabis,
though less well-expressed n that genus. The posterior lobe of Pa
and P4 is divided into two distinct cusps: a broad labial hypoconid
and a narrow, less pronounced lingual entoconid. The entoconid
attains much greater development in P4, so that with continued wear
the entoconid and hypoconid join posteriorly to form a posterior lake,
in Pa the entoconid is so weakly expressed that'at no stage of wear
does a posterior lake form This condition obtams also in Protolabis
Ps tends to be rectangular, the anterior and posterior widths being
approximately the same, whereas P, has the wedge shape characteris-
tic of most camels. The molar teeth are high-crowned and very simple.
In P. grandis the stylids are much more prominent
The body of the rams is slender and graceful, its depth being
not so great as in Oxydactylus or so variable anteroposteriorly as in
Pliatchenia. A large mental foramen occurs below and anterior to
Pi, a second, much smaller foramen is situated directly below the
posterior edge of P4. The dorsal border of the mandibular foramen
is approximately level with the occlusal surface of M,. In contrast to
Oxydactylus and Nothotylopus, there is no downward expansion of
the posteroinferior border of the mandible, the ascending ramus
makes a steeper angle with the body of the mandible in this and
other species of Procamelus than in Oxydactylu. The angular process
is quite prominent and is canted lngually. The condyloid process re-
sembles that of Oxydactylu. The long, slender coronoid process of
P. occidentais specimens from other areas extends well back over
the condyle, in marked contrast to the short, stubby coronoid m
Nothotylopus.
Metacarpals of P. occidentals average 336.3 mm in length, meta-
tarsals330.0 mm. The average length of the radius-ulnae, exclusive of
the olecranon, is 368 mm.

Procamelus grandis Gregory, 1942
Table 13
TYPE. UCMP 32864; skull and jaws.
Occ Eca. Lapara Creek Fauna.
REER MATRIL. UTBEG 31081429, 3086-531.
DEScmNN. The identification of P. grandis is based on two






Table 18. MtmhanmEN or Procamaeu grande
Fence Line Loc Big Spring Canyon Bee County Bee County
Mandble UCMP 83473 UCMP 8301 UTBEG 31081429 UTBE 80896-531

Distance betwn igl and cemlye 34.0 450 28.0
. P" of Jaw posteror end of Mt 69.1 640 64.0 60.0
Depth of faw at anterior end o M 47.9 87.0 430 8.0
Depth below /C 28. 20.4
/C-P, 21.4 19.0
PI-P. 19.1 294
P*P, 43.9 41.0 45.0' 1050
M,-MI 96.8 1020 104 0
P-M. 137.8 144.0 1500'
C length x width 132 x 9.6
P,, length x width 10A x 7.0
P. length x width 13.1 x 63
P,, length x width 13.5 X 82 14.5 X 58
P, length x th width 1.9 10.5 202 8.8
,M length X width 2.9 x 17.1 14.3 266 X 160
M, length x width 29.3 x 18.9 0.8 x 14.9 840 16.5
Mh. length x width 45.8 X 19.2 458 X 14.9 46.0 x 166
'1A Webb <(1% ftar GT (IU2)
'Aft Webb (109)








partial lower jaws: UTBEG 31081-429, a left mandible possessing a
broken Mi and Ms-Ma and alveoli for P-P4, and a virtually complete
ascending rams, and UTBEG 30896-531, a right mandible with
P4-Ma. In general morphology the Lapara Creek specimens closely
resemble the holotype of P. grandis described by Gregory (1942) and
P. gmndis specimens from the Burge Fauna figured by Webb (1969).
With the exception of a single P4 preserved on UTBEG 31081-429,
the Texas specimens lack the diagnostically valuable premolars. Some
minor differences in the molars and in mandible configuration are
observable, but whether or not these difference are geographically
consistent, or even diagnostically rehable, is uncertain. If the man-
dible (UCMP 33473) from the Fence Line Localty (Webb, 1969)
is representative of the Burge P. grand, then the lower molars of
members of that population appear to be shorter and wider than
those from the Lapara Creek Fauna (Table 13). The molars of both
the Burge and Lapara Creek specimens are narrower than those of
P. robustus. No important differences between the Burge and Lapara
Creek specimens are discernible in characters of PA. Although the tips
of the coronoid processes of the holotype (UCMP 32864) and para-
type (UCMP 32589) of P. grandis from Big Spring Canyon and of
UTBEG 31081-429 are broken off, they appear to lack the pro-
nounced posterior sweep of the Burge specimen (UCMP 32286).
Other minor differences, and perhaps those listed above as well,
are apparently attributable to normal individual variation. On the
















Founs 1t7Not ftylopus camptognaths gen et sp (UTBEG 31081-20,
'Fgy); let ramu, labial view IW-M, oclusal view PrM.








basis of presently inadequate material there seems to be no way to
separate or recognize geographic variants in this species.
Because nothing is definitely known of the metapodials of the
new Lapara Creek genus, Nothoylopus, the large (average length =
460 mm) Procamels-hke metapodials from the same Lapara Creek
quarry from which both P grandis and Nothotylopus were recovered
are not confidently assignable to either taxon

Nothotylopus new genus
A virtually complete mandible of a medium-sized camel from
the Lapara Creek Fauna of Bee County, Texas, is believed to repre-
sent a lineage distinct from previously recorded extinct camels and
whose origins may be found within the Protolabis stock. Until further
material is forthcoming, generic characters will correspond to those
of the type species, Nothtylopus camptognthus.

Nothotylopus camptognathus new species
Figures 17, 18, Table 14
T-PE. UTBEG 31081-26, a left mandible with I,-M0.
TYPE LocArY. Farish Ranch, Site 15, Bee County, Texas
OcCU RENE. Lapara Creek Fauna.
ETYMOLOTy. nothos Gr. spurious, bastard, n reference to its
uncertain origins; tyiopus = Gr camel; kamptos Gr. curved, bent;
gnothos = Gr. jaw.
Dr AnosIs. A medium-sized camel with widely flared and curved
masseteric region; short, weak coronoid process; long Pi-Ps diastema;
Ps absent; lower premolars anteroposteriorly compressed but complex;
molars robust and low-crowned with pronounced anterolabial ribs
DESCRPrION. Identification and systematic placement of this
species is based on a well-preserved left mandible with complete
dentition. Dental formula: S1CIPaMs Shape and proportions of the
lower jaw of N. camptognathes are very distinctive. The masseteric
region is widely expanded and broadly curved labially (Figs. 17, 18).
The prominnnt angular process occurs high on the ascending ramus
well above the occlusal level and lacks the strong inflection seen in
Procamelus. The condylar process is thick and broad. In contrast to
the broader, backsweeping coronoid process 'of Procamelus and
Pliachena, the coronoid process of N camptognathus is thin, nar-
row, and exteniJ vertically to a level only slightly above that of the
condyle The maieferic fossa is deep and ovate. On the lingual side






PAMON TEXAS AATIODACIVI


of the ascending ramus the large mandibular foramen occurs at a level
approximately halfway between that of the oazlusal surface and the
angular process. The inferior border of the body of the ramus rises
obliquely with respect to the superior border until just anterior to Pa it
becomes parallel to the horizontal axis of the mandible The ramus
deepens again at the posterior end of the symphysis, forming a lump
on the inferior border at that point. There is a large mental foramen
below the anterior border of PL and a small one below Ps. From a
point just anterior to Pi the symphsis widens towards the anterior tip.
This widening is further emphasized by the outward flare of the lat-
eral incisors. A deep groove about 15 mm long occurs on the inferior
side of the symphysis along the symphysial suture between the first
lower ncisors.
The incisors are procumbent and slightly spatulate A short, 9 mm
diastema occurs between Is and /C. The canine is strong and slightly
recured, but lacks the sharp anterior blade present in rocaelus.
Pt is small, two-rooted, and only mildly caninifom, differing in each
respect from P of Proamelus. P is absent The diastema between
PI and Pa is more than twice as long as that between /C and Pi. Pa
and P4 are more compressed anteroposteriorly than in Procamelus or
Pliauchnia. Although small, Ps has three well-developed lobes with
easily discernible minor cusps, a condition contrasting with that of
Procamelus, Pliauchenia, or Protolabis. It is also shorter crowned than
Procamelus or Plauchenia. The lobes are separated by two small
labial folds and a more pronounced anterolingual flexid. The posterior
lobe is composed of a major labial cusp (hypoconid?) and a minor
lingual one (entoconid?). Continued wear should result in the for-
mation of a small posterior lake between these cusps. The middle lobe
(metastylid?) is high but blunt The anterior lobe (paraconid?) is
flexed anterolingually a and exhibits a sight anterior bifurcation is
charaetnstically wedge-shaped, but is shorter than in other contem-
poraneous camels. A deep posterolabial fexid lies between the middle
and posterior lobes and a wider, more prominent one between the
middle and anterior lobes. Smaller folds occur anterolabially and pos-
terolingually A distinct posterior lake has formed between entoconid
and hypoconid. The lower molars of N. camptognathus are relatively
wider and less hypsodont than those of Procarelus and Phaschenia
(Tables 12-14). Ma and M. have well-dveloped anterolabial ribs,
a character that may be found in some other camels, e.g., Miolahbs,
although it.does not appear in Protoabis, Procamels, or Pliuchenia.
Extreme wiar'prevents the determination of this character on Mi. A









Table 14. MEasU~nMrrs or THE TyrF OF Nothoylopws campotgnathus
Lower law UTBEG 31081-26
Height vertical ramus 186 0
Condyle to angular process 315
Condyle to coronoid process 21 0
Depth of ramus below center of M 60 7
Depth below center of Mi 497
Depth below center of Pt-P diastema 35.8
Length P.-Pi diastema 615
Length Pi-C distema 28 1
P.-M. 184,0
PrM 1084
PI-M 977
P.-M 24
M-M, 84 4
P., length X width 118 x 58
P., length x width 138 x 85
M,, length X width 19.2 x 14 0
MI, length X width 274 x 165
Ms, length x width 397 x 180
'Approxlmain

tiny accessory tubercle is visible at the base of the crown between
the anterior and posterior crescents of Mi and Ms. This feature is not
characteristic of the other advanced camels with which Nothotylopus
is contemporary.
DiscUSSION. In view of the incomplete material available for
comparison, derivation of Nothotylopus from any of the currently
known groups of camels is difcult and precise systematic placement
tenuous. Nothotylopus exhibits a strange mixture of advanced and
conservative features: long diastemata, reduced but complex Ps and
P4; P2 absent but P1 two-rooted; and robust, buttressed molars
Reduction and loss of lower premolars has been one of the most
consistently employed and reliable characters in differentiating Late
Tertiary camels. With the possible exception of Aepycameaus (= Ali-
camelus), all of the major taxa, i.e. Protolabis, Procamelus, Pliau-
chenia, Megatylopus, and Miolabis exhibit some degree of premolar
reduction, including the loss of Ps and/or Pa. Heretofore loss of Ps
has been a diagnostic character restrated, at least among the large
Tertiary camels, to Plianchenia (Cope, 1874a) and Megatylopus
(Matthew and Cook, 1909) although individuals in some species of
Protolabis, e.g. P. longiceps, may lose P2 Whereas reduction may be
a parallel convergent character acquired separately by each of these
genera, it is more likely a progressive character peculiar to the Proto-








labis lineage. For example, there has been reduction and occasional
loss of P2 in Protolabis, reduction in Procamelus, and loss in Phau-
chenia and Megatylopus. Nothotylopus possesses this and other, sep-
arately acquired, progressive characters, but retains enough conserv-
ative features to preclude its derivation from any but the Protolabis
stock. This conclusion is based primarily on short, two-rooted P, and
reduced Pi and P4 of Protolabis. These features are also present n
the small gracitls group of procamelines and in Mwolabs However,
the former is a separate, early offshoot of the procamehnes, whereas
Miolabis, as it is now known, is taxonomically uncertain, perhaps
ultimately to be placed in synonymy with Protolabis. The Oxydac
tylus, Aepycamelus, and Hesperocamehls lineage is far too conserv-
ative in premolar reduction, and too advanced in upper incisor reduc-
tion and dimensions of skull to provide a possible direct ancestor for
Nothotylopus, while the Procamelus-Plhauchena-Megatylopus groups
are too specialized, especially in degree of hypsodonty The probable
ancestral stock, therefore, should be found among the early members
of the genus Protolabis, or perhaps some form intermediate between
Protolabis and that line of oxydactylines which presumably gave rise
to it















rea




FIcunR 18, Nothotylpus camptogathus n. gen et sp (UTBEG 31081-2,
Type), left rams, labial and oceusal views









Megatylopus Matthew and Cook, 1909
In 1875 Cope proposed the genus Pliauchenia to include those
camels lacking the second premolar of the lower jaw. In addition to
the type species, P. humpreystana, he described another species, P
vulcainrum, from an upper dentition in which P2 was present The
dental formula for the new genus was therefore given as premolarss
4" P. vulcanorm has subsequently been assigned by Gregory (1942)
to Procamelus on the basis of retained P2, but lack of Tu has been
used as a diagnostic character for all later additions to the genus. In-
cluded in this taxon are various presumably related forms which,
while sharing P loss and succeedig premolar reduction, are clearly
distinct on other grounds Therefore, as Matthew (unpubl. MS) sug-
gested, it appears that P2 loss marks a stage in the evolution of the
protolabine camels and is not characteristic of a group of nearly
related species In an attempt to rectify part of this confusion,
Matthew and Cook (1909) restricted the very large species of Pli-
auchenia to a separate subgenus, Megatylopus, which has since been
elevated to generic rank (Gregory, 1937).



I










FGlunr 19. Megatylopus pnnmevus n sp. (UTBEG 31081-460, Type) left
ramus, labial and ocelusal views P,-M,

Matthew and Cook (1909) originally described Megatylopus as
having the following characteristics
Gigantic camels with denition I C' Peaf M|, the second premolar absent in
both jaws, the first retarded or absent, the reduction of the posterior premnolar
intermediate between Procamelus and Camelus Upper molars with prominent
external nbs and styles, as m Auchenia Limbs and feet moderately elongate aid
very massive, large m proportion to size of skull








Although additional knowledge of Megatylopus has accumulated
since its original description, the genus is still imperfectly understood
and, therefore, an comp and/or systematic conclusicomparsons ad/or systmati conclusions re-
garding it and related forms are necessarily tentative.
Species that have been included in Megatylopus are the type spe-
cies, M. gigas Matthew and Cook (1909), M spatula (Cope, 1893),
M. merriami (Frick, 1921), M. maior (Leidy, 1886) and M. cochrani
(Hibbard and Riggs, 1949). M spatula and M. errtnmi are regarded
by Webb (1965) to be properly placed i Titanotylopus Barbour
and Schlutz (1934). Webb (1965) bases his conclusion in part, on
the lack of a cranial flexure, the broad anterior lobe of P(, and the non-
reduced Pa, features that these two species share in common with Ti-
tanotylopus, but which serve to distinguish them from Megatylopus
To the remaining species of Megatylopus must be added two new
forms, M matthews from the Hemphillian Coffee Ranch Quarry, a
distinct species which Matthew (unpubL MS) recognized as Para-
camelus arencola, which is listed but not described by Reed and
Longnecker (1932) and which Webb (1965) described and a rather
primitive form from the Lapara Creek Fauna, M. primaeous, which
is described below.

Megatylopus primaoeus new species
Figures 19, 20, Table 15
TYPE UTBEG 31081-460, a left mandible, with P2 alveolus,
Ps-M$.
TYPE LOCALT. Farish Ranch, Bee County, Texas.
OCCUmRENCE. Lapara Creek Fauna
REFERRED MATEInla. UTBEG 31081-18A, a partial right ramus
with Ps-Ma; 31081-18B, a metapodial.
EnTYMOLOc prtmaevus = L early, young.
DIAGraNO s. An early species of Megatylopus, similar to M.
major, but with a small PF and a larger and more complex FP, man-
dible slender and vertically shallow.
DESCRIPTION. The type mandible of M primaeous is long and
slender, the depth of the ramus being consistently shallower than
Nothotylopus or Protolabis (Tables 11, 14, 15) Characters of the
ascending ramus are mostly indeterminate because of poor preserva-
tion A small mental foramen occurs just below the center of the an
terior half of Mi, the anterior mental foramen is situated above the
posteronferior border of the symphysis where it deepens ventrally.
The alveolus anterior to this foramen contained the lower canine.








Above the foramen and just below the surface of the bone is the un-
erupted Pi. Two small alveoli for P2 occur anterior to Ps This tooth
was obviously quite small and probably nonfunctional. In view of the
fact that the more advanced species of Megatylopus are characterized
by loss of P2, the retenton of a small P2 m M. prnmaecus is regarded
as representing the last stage of reduction of that tooth in the Mega-
tylopus lineage
The diastena between /C and Pa is approximately 62 mm long
Pa is two-rooted and has well-developed anteror and posterior cus-
pids The central cuspid (metastylid) is unworn and attains a height
of 18 mm above the base of the crown. The crown flares from the
base to reach its greatest anteroposterior length (178 mm) at about
two-thirds its vertical height The anterior lobe or cuspid is rather
deeply grooved but only weakly inflected lingually The posterior
lobe has a large posterolabial cuspid (hypoconid) that is strongly m-
flected lingually, its lower slope extending farther lingually than the
weakly expressed posterolingual cuspid, or entoconid. These two
cuspids do not join towards the base, thus continued wear would not
produce a posterior lake.
P4 is a much larger and more robust tooth than P8, and is quite
hypsodont. The maximum height of the tooth, only slightly worn,
is about 27 mm. The anterior cuspid is lingually iflected and set off
from the primary cuspid by a promient groove. A faint posterolabial
groove is present between the principal and two posterior cuspids.
The posterior cuspids are equally prominent and join posteriorly In
medium stages of wear they will form a posterior lake.

Table 15. MeASUREMENTS or Megatylopus prnmaevus
UTBEG
Lower law 31081460 (Type)
Condyle to anlar process 7
Depth of jaw blow M 50
Depth of law below M 33W 0
Depth of law below P-P, diastema 30,0
prM, I1590
PM, 142 0
pR-P 89 0
M,-M, 1200
P., length x width 178 x 8.2
P., length x width 222 12.0
M,, length X width 84 0 x 18,8
M., length x width 430 x 21.0
Me length x width So8 X 200








The lower molars are hypsodont and are characterized by strongly
developed styhds. Mesostylid and metastylid are especially prominent
in the holotype In the only other ramus referred to M. primenus
(UTBEG 31081-18A), these stylids are virtually absent. This jaw
differs also in its smaller size and more reduced premolars. Whether
or not the differences between the two specimens now assigned to
M. primwevus are significant is for the present indeterminate.
A large, massive metapodial (UTBEG 31081-18B) is tentatively
assigned to this species.
DISCUSION. Of the species of Megatylopus accepted in this
paper (M. gigas, M. maor, M. matthewi, M. cochrani, and M. pri
maerus), M. primawous is the most primitive form yet recognized
In a clearly discernible evolutionary trend towards the condition
reached by the genus Camelops (Webb, 1965), the premolars show
a continuous reduction in size and number. P2 having already been
lost in all species but M. primoaeus, this involves the reduction and
eventual loss of P, and Pa. The absence of Pa (and the usual absence
of Pi) represents the taxonomic line of demarcation between Megaty-
lopus and Camelops. M. cochranm from the Rexroad Fauna of Kansas
has been demonstrated by Hibbard and Riggs (1949) to be very
similar to Camelops and probably represents the most advanced
species of the Megatylopus lineage. Unaware of the Lapara Creek
specimens, Webb (1965) considered M. maotr from Mixson's Bone
Bed in Florida to occupy the earliest level of Megatylopus evolution
and placed the species M. matthewi from the Hemphillian of Texas
between those stages represented by M. gsgas and M. cochrani. This
arrangement is based primarily on the condition of Po In all species
but M major and M. prmaevus, the roots of Pa are fused. These
species also exhibit greater reduction in size and complexity of Pa
than is attained by M. major and M. primaevus. On these grounds,
M. primaeous and M. major are clearly more primitive than M.
matthewi and the two succeeding species. M. major and M. primaecus
are similar and probably closely related, but because of the retention
of a small P2 and the greater complexity and size of Ps in M. primae-
tus, I consider it to be a more primitive species than M. major The
differences are not great, but because the characters employed in
taxanomic differentiation of these forms also reflect relative advances
or stages in their evolution, it is useful to recognize and emphasize
such distinctions. This is most apparent when considering these
species from the standpoint of their stratigraphic occurrence. The
evolutionary series beginning with M. primaevus and continuing


























fl mrE 20 Megatyopun primaeous n sp (UTBEG 3108100, Type), left
ramus, labial and occlsal views

through M. major, M. matthevwi M. gigas, and M. cochrans cor-
responds with their relative stratigraphic position. Complcations in
this evolutionary-stratigraphic sequence arising from different rates
of evolution in different geographic regions, intermrgrations, etc,
are not as yet apparent


Subfamily Floridairagulinae Maglio, 1966
Floritdaragulus White, 1940
The exact systematic position of these puzzling artiodactyls is
uncertain. The fragmentary nature of flordatraguline material from
Texas and Florida deposits has prevented much speculation on
phyletic ainities, but it is hoped that study of skull and jaw ma-
terial now available from the Thomas Farm will lead to a better
understanding of the group. As I have previously commented (Pat-
ton, 1966(67): 184).
Floridatragtlss bears considerable resemblance to members of two family,
of artiodactyls, the Hypertraguldae and the Camnehdac Flondatragulus resembles
members of the Hypertragubdae in the presence of the ntercolumnar pillars, a
double enamel loop of the heel of MW, rather prominent eingula, and in the
occurrence of a diastema between P. and P[ In the hlypertraguhds the detached
P, tends to be unicuspid, in Floridatragulus this tooth is bicuspid, though weakly
so The anterior lower premolars of Flondairagulus are laterally compressed as in
cmels, and P,, although more foreshortlned than in most amels, retains the








general features of that group In the skull and jaws of Flondatragula from the
Thomas Farm, now in the Frick Laboratory, the extremely elongated muzzle has
four canimform teeth (or alveoh) on each side, probably representing modifica-
trans of the mncsors, C/, and '. This is a characterstic feature of the Tylopoda,
in which the upper incisors and anterior premolars may undergo reduction, but
in which there is never a complete loss of the upper inisors In contrast, the
hypertragulids are marked by ether extreme reduction or complete suppression
of the upper incisors, although they may be retained in some of the very pramitve
types.
Because of its overall resemblance to the camels, in the same
paper I referred Floridatragulus to the Camelidae as a new sub-
family'. I felt that those features that distinguish it from the Cam-
elinae are certainly equal in taxonomic weight to those of the other
camel subfamilies Apparently for similar reasons, White (1947)
tentatively placed Nothokemas as the type genus of a new family in
the Hypertraguloidea Nothokemas' affinites with the Camelinae are
rather substantially documented, but similar assurance cannot be
claimed for Floridatragulus.
Clearly while Floridatragulus has several distinctive camelid
features such as the "hook" on the angle of the lower jaw, the canmi-
form condition of the upper anterior dentition, and the relatively
swollen bullae, other traits are characteristically "ruminant" (fide
Simpson, 1945) For example the enclosed orbit, the presence on the
molars of intercolumnar tubercles, cingula, strong ribs and styles, and
the double lobe on the hypoconuihd of Ms are all quite primitive
features not fully shared by any of the camels, especially the later
ones That the similarity between the various artiodactyl group
increases back through geologic time is indisputable (Matthew, 1905,
Scott, 1940, Colbert, 1941, Simpson, 1945). However, although the
camels and ruminants are supposedly recognizable as dastmct groups
as early as the late Eocene, Floridatragulus shares relatively few
features in common with contemporary representatives of its basal
stock. Whether or not the morphological similarity between it and
the Camelidae Is the result of direct evolution, parallelism or con-
vergence is not immediately determmable. For the present, available
evidence suggests that Floridatrogulus descends from an early and
distinct offshoot of the basal tylopod stock and that it (and Notho-
kemas) subsequently occupied an adaptive zone in the Coastal Plain
similar to that occupied by some camels in the Great Plains. With
the invasion of the Gulf Coast by several diversified artodactyl genera
A similar conclusonn was reached by Maghia (1966) as the quoted paper was
m press








(including other camels) during the late Miocene and early Pliocene,
the autochthonous Gulf Coast genera became extinct
The known species of Floridatragulus, so far restricted to the Gulf
Coastal Plain Miocene, are separable into three size groups that cor-
respond with their stratigraphic occurrence The smallest group is
represented by F. nanus n. sp from the Garvin Gully and F barbourn
known only from Thomas Farm. The medium-sized F. texanus n sp
and F. dolichanthereus have been recovered from the Burkeville
Fauna of Texas and the uppermost strata of the Thomas Farm quarry,
respectively. The stratigraphically higher Cold Spring Fauna of
Texas has yielded a still larger form, F. hesperus n. sp., which is
12 per cent larger than F. dolichanthereus and 20 per cent larger
than F barbouri













Icm
FIGmE 21. Floor latragulu nanus n sp. (UTBEG 40067-i94, Type), left M,
occlusal and labial views

Floridiaragulus nanus new species
Figure 21
TYPE. UTBEC 40067-194, a left Ms.
TYPE Locarr Hidalgo Bluff, Washington County, Texas.
OCCURaRECE Garvin Gully Fauna.
ETroro.oc. nanos Gr. small, dwarf
DIAiNoss. The smallest known species of Floridatragulus, m-
tercolumnar tubercle strongly developed between protoconid and
hypoconid of Ms, weakly developed between hypoconid and hypoco-
nulid, talonid of Ma divided, but differs from that of other species of
Floridatragulus in having a posterior cngulum and two small tuber-
cular cusps developed posteriorly between the two grinding surfaces.








DESCRIaTON. This species is recognized i the Garvin Gully
Fauna by a single tooth collected from the Oakville Formation ex-
posed at Hidalgo Bluff, Washington County, Texas. This specimen
differs from those representing later occurring species of Florida-
tragulus not only in its small size (20.0 mm X 10.0 mm vs. 24 mm
X 12 mm for F. barhouri), but in the nature of the talonid of Ma
Whereas all known species of Floridatragulus have an invaginated
talomd on Ma, none but F. nanus exhibits any cusp development
between the two resultant grinding surfaces. Rising between the di-
vision of the talonid at its posterior border are two small icspules, ap-
parently developed from mfoldings of the posterior wall of the
hypoconulid. Both labial and lingual cuspules show wear. In addi-
tion to the above mentioned minor cusps, a small but distinct cin-
gulum appears on the talonid of Ms, a feature also unique to this
species of Floridatragduls.

Floridatragulus texanus new species
Figures 22, 24B, 24C, Table 16
TYPE. UTBEG 31190-28, a pair of lower jaws with right P2-Ma
and left Pa-Pa and Mi-Ma.
TYrP LOCxALr. Near Point Blank, San Jacinto County, Texas.
OOCURENCE. Burkeville Fauna.
DIAGNOSIs. A medium-sized artiodactyl with brachyodont molars
and a very long mandibular symphysial region; diastema between
P2 and Pa equal to two thirds of that between Pi and P2, intercol-
umnar tubercles occur between protoconid and hypoconid in Mi and
Ms; talonid of Ma divided into two separate grinding surfaces. Differs
from F. dolichanthereus in having larger premolars, and in the less
invaginated talonid of Ms.
DESCRPnON. Although the lower molars of Floidatragulus re-
semble those of Prosynthetoceas, they are separable from that genus
on the basis of lower crown height, a larger and divided hypoconulid
on M8, and the presence of an intercolumnar tubercle between the
protoconid and hypoconid on Ms. The lower premolars are generally
narrower and less reduced anteroposteriorly than those of Prosyn-
thetoceras. When the complete mandible and dentition are preserved,
the two genera are easily separated by the presence of diastemata
separating tde anterior premolars of Floridatragulus.
Discussir. The Texas specimens of F. texanim are very closely
similar to the one White (1940) described from the Thomas Farm




















Icm
FloRa 22. Floridadrgulu texanus n sp (UTBEG 31190-28, Type), left
rams, labia andal and oual viws PM

deposits of Florida as F. dolichantheres. The Burkeville specimens,
not known to White, differ from the Florida type specimen (MCZ
3035) and material subsequently assigned to the same taxon in the
slightly larger premolars and in the lesser development of the division
of the talonid of Ms, a feature White emphasized in the original
description. In the Thomas Farm specimen the invagination of the
posterior border of the talonid of Ma, a reflection of this division, is
pronounced and extends from the oecdusal surface down to the dorsal
edge of the basal cingouum. The Burkeville specimens, which exhibit
approximately the same amount of wear, show only a very slight in-
folding of the posterior border of the talonid, which extends no
farther down than 1.0 mm from the occlusal surface. Although the
Texas and Florida specimens are very similar, the above-mentioned
differences may be real and may possibly reflect a minor genetic
divergence resulting from geographic isolation of indefinite, but
probably amall extent and duration. If these differences prove to
be consistent, then they assume not only taxononuc, but also zoogeo-
graphic importance. For these reasons, I feel that they warrant specific
separation.
Flokdatragulus barbouri White differs from F. dolichanthereus in
being slightly smaller and in the shorter diastema between Pa and P2.
The anterior half, metaconid and protoconid, of Mi of the holotype
(MCZ 4086) of F. barbouri is extremely foreshortened, but judgmg
from the way it is crowded between Ma and P4, this is probably a
pathologic condition. No such foreshortening appears in any other
I








of the known specimens of Floridatragulus. Whether F. dlichan-
theresn and F. barbouri lived contemporaneously, or whether they
represent an actual ancestor-descendent lineage is uncertain The
nature of the Thomas Farm deposit and the history of its excavation
(White, 1942; Bader, 1958) suggest that those forms as they are
known so far were separated temporally. The largest species, F.
dolichantherews, was recovered from the uppermost layers of the
deposit and has not been found in any of the deeper sediments
(White, 1942: 30) The smaller, less advanced F. barbourt was re-
covered from a deeper portion of the quarry. Hence there is a reason
to accept the second alternative, that F. barbouri represents an earlier
group that evolved (no evidence suggests replacement) into F dolf-
chanthereus. Such an interpretation has obvious chronologic .mpli-
cations (see Age and Correlation).

Table 16. MusmsasuMrs or Floodatragulus dolichanthereu At texanus n sp.

MCZ 835 UTBEG 31190-28
Lowerjaw (TypefF do) (TypeofF. tex )
Length P-P, diastema 33.4
Length P-P diastema 295
PrMc 70.7
Ps-P 22.7
M-M. 55 0
P., length x wdth 12.1 X 3.9
Pi, length x width 110 x 49
P., length x width 11.0 x 5
M,, length x width 130 x 8.5 132 x 91
M,, length x width 163 x 11.0 18.1 X 11.4
M., length X width 256 x 12.6 25.2 X 12.0


Florkdatragulus hesperus new species
Figures 23, 24a, Table 17
TYPe UTBEG 31219-266, an incomplete right mandible with
MI-Ms.
TYPs, LOCATr Donahoe Farm, near Cold Spring, San Jacinto
County, Texas.
OcCURRENCE, Cold Spring Fauna
ErTYMOLOy. hesperos = Cr. west.
DIAcNOs~. A large species of Floridatraglus, some 12 per cent
larger than,. dolichanthereus. It is characterized by a rather large,
recurved mesastylid on Ms, and an intercolumnar .tubercle between







I.BULL.E .IN FLO.UHlA S IAlE 1 MLUSEUMI.


Icm


FIC 'PF 2.3., Flori daltraigtlu, s h .spt-rs ii. sp. I7 1 'l IEG .31.2 I -2 6, '.'p r. iht
2r31mus, labial and UO _,_uISl vi

'Talet 17. MEA.SL'-REMENTS OF Finriddfraiii._:. hq., 'eru;:


STBEG
31219-266 (Tvp


Lower jaw


-11,-Mu
M:. ingth d width
;.:, ] .te 'l th ". \v'ilth
\,^ lenigth :. widtii


6:.0'
15.1 11.0
19.2 I 12.i
29. -. -" 12.^'


1 r T-p. r i I fat.C-

the ih.poconid and hypoconilid inl ;lddit](ionJ to tIhe on.l be'tw'ln thi
hypocn1jdt ard pro)tocoWtid of M;,.
DISi.T SSit(1 N. This largrn^r forI! ftl11 tlile r O- S[)ri Fa i 1mina Co-
curI, highest str.it i)r;ipi. i a llv of o ll the 11 F.urhid tra ulr u.v spcci's a.,nd
rpres.rsc lts the moist advaInce'd sat h hn .2 s reaichi-'d. In :c ld itiorn
to its ]arT.' 17 the pIre.l .' tc .. f tit intl:.rc luol rni3ar tiT.iberele or, tuh
i1alnir of N.:1, the princiiunced r-ie0.stvlid, aind the better develop-d
anterior cingul -iir l .I s: erv,' to dlistiirlai tlI it.

Suborder RH uII. Ai N A Sc.opoli. 17 77
Ilnf 1 ora cr T 'I.-) \ icl..I F ,\ I 'c r. ] .8 5;}
FKinilv P1 iOTOC.ERATIDAL' MIarsh. 1S91I
SNbfiilamiily SYNTIIFTrCT:iIATiN. ric -'k, j~ l 37
The pe'(cliarly horned svnthtltocerines coi prisre the iilost a.ll--
(dantiv represeri)Tttd ru'Op of artiudactvls Ircover.d .ro in the 1 icic'tnc


Vol., 14,


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"1F sviK n VllymV f rfrI n, l( / ",! O'( TNs ptcie ha- s 1b e-n l itreaed -r 1..vE111 ,i,,
(Pt Itoin, 1966: 67 1967 7
Since ',-..ti t 1T. 1 tii]. m .i s i ::rt.t Ip fiad tlt. (vppI rtiunl it'v ft
b)'iln a n3 .v sii. v, it- l B.-' TI.avior of tIh A I ri ..c\L.i Lu.f..1n of(P
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Il''rik ( il?-P ti. i4 Ii \\ }ji. i not LlAv4iilalil ,tt 1111r tililt this x'ai> \\3fittwH.


Pr, 's./ r.f t. to"'rs [rick 1937
i(({FCJ.\ /utifyxuuu.\ I f
Figures 2.. 26. 27, l"ihs S 19
/..:,n m' ". ry. tA'. u'. il..', -124. p. le. pl. II. Fi s. S-12
D n:??r iF? 'tv rnflrWitiei' }l.Ity., 1424. p. 1@, pi. 1I l r). 7.
?A(raJ J .I s rri% IJ, I I 2-, ` i I 7. pl, I ? fi, 1i
SPr..t,, i,., fr.,,,.?i.,, i tiiv 1 -24. p. 1 1, l?1. III. fiL.^ .-"
-.i .11i/ .Is. sp (i,. Si, pe'.o :, ..;', p .37.
?C ra' t..- ,,. ',' ', f. ., s,.. L a Frid-., I'.i'7 p. "2. ':7
?S /, ,f et;," :-T., r.;.; ri !i !':k, 1 37, p I:0,3. -, fli f
B Iya.StIrf .r(T t in.O l \V IAi W oo a Id j\V I !7 p 1:.!. p I, tu ':- 5, 6

Ir' i.L r/ t.x nr II IC:. 3 -12. p 1. .' L 'Sr 'n .-jra( frEx!?if. r. p. 1C 7.
fip A.c 'i
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(T cw f ilrlc-r. l(aTrvift (I.iliv i'r. : l'rkU.vilic I, 1Ai; .
i-' FkH.RED NMATFRIAi.. I'T[BE 3 j114-j1 iirnxia with P -N ,
TAM. U uinirnrih., pmart-fi right ijnaxi.ll -With al]\olht.s for 1 -'. and P -MN
UTDEC 31190-31, p i.il l 31. 190s 75, partial ro.tral horn rr, i-'
I.1S-S1, aritu-ior porti.,n ot left rarrnus with PMNi1. 31190.-9, left
P.,-\. 31190- 02, l. \1,-\.., S\11. ..S 1,ft P.,-M. anrd numerous
isolated IeIth.
DiA..Cesvs. A reM it i0 .sinalS N1 U w t!ir ilnti vit1t ;h dlsh-Sin hcl
no.s :il. r lOa r and-il u)1 i pIsut I( nl t h .'(t i- r ftLi11,'dt. LUpptr L I :i( r- a .1 ra '
.hort. broad, aii j( NI Ti \ t p,.rT.,rl ti \aird oc) cI.1]s l s rf'accf.', th '' .ire
(ch.rterrizt piOrini eTit meAdi'd V I \'- )Iap'd pil :.irs anid I 'i -delike
C'I0 ula. a.nte ri.v 1(d pI .t-riovi. Pl' r'] i' a pri 'z i ITed develop
rilec-it o.if Sty'Its, t-i'sI)e dIv tht. rr,.i o.s.' P! p9ns i, t. 1;: is redt.iced. !)ut
.V o-rooted.


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-1 vp ii )I -I d 0 1 ( ti. 1 142t (3. !:I
The lower preriolda.rs are al reduced, esprcia l I' i,. P; is. lo-
rooted and is similar t.o but .siialle'r than P3. Ti] p)rimitar\ dijEt-renC.s
I It'f.';cn Z :L .11id. I:. i ti -I "Sr ItT- sI'.\p ;V4. io.I of the ~ I t .li] iJ o ll ..tI

fitter P., is el: I te i d i .t A d ligt 1tl V I)roitdler posterii urv thiTa
aiiteritirl,. I'h.t h-I (poE ild L.nd protoconid arc i ore;' distinct Eri'f'll
tit" ilc't c.ci.nid tii.;tii ill I .As in P- sW. clIints at 'he has- ,c i th,
htp.c"rliC id proloconid ari nioticcable. Pi is slightly wede-shaped.
N.lost of the redluti'on in this tooth has .,occLr-edl ] in the rI'ton of
the protocond, ii. 1le nu^ticoild is hi i and 11slC rp t, ('ite .lntoOJCfuid is
1 ,i O v i.id 1l.,.lik e. TLh iry )oc i'o Es l ext ri,. p- stelEi r[\' .:,one ILs
0. i: y to |o i.) (I i i c:-ji'tOc'Oni (I 1) wo:>rii spfci ij sI tle rnel ,'if ,Iic],
einloc.if .d d I % l i't"-T-li ] IT l o fiti T ;hll i : 1i T:. l 'ii il i is 1 't. "f i
a: se l-. h., y :poe.tiidi is ,.(v l i 3toI t -ii .t h1 -,i ;t t I i. post'o.t i-r l.i, -t I
t"d.(I of ti t()t.li. Ihitrc ai -ss(. distint I c w lli L ith' j-.base of
ithe protocoilit .

Ihe lovw'r rm1 oas of P. t.vanins.s r-I- .siibhyv;psodo-nt. The ereseernts
i. prticonlid, hypocoiid arc syalnelrical Ind shlrCI ow little tenidcilcy
to..virtl.s r.cui .in.in Sty lids are reduced iin compariso-ln with thosi,.
OHn th' uipper n'iolar]i; ort tl l riet: tI lid is prTt"imolinced. The anterior
c'r, -s.' ln i( met a c, o ii oV'r ].ips t- i pi). s-.rir (n r cenit i( eil toC lid
.1ii1htI tL fu t i. t.Ail it t ;. i t Ih A I 1a I I '( d 1 1i] p 11ir o( ccurs at th
: is of tin-e cr v I .lt '$ ll' tw Ili i I t r pl o to.toriid t ,I il I f. ri(l3Vi f
1 h.poco i .1 i (T sts o NI,. 1 lir)t rust to th. 1(iU ll 1il ;TS ,1i !t i' lIpperT Trila s, thi I V.,o T
(sit.> \VWOld l ot i Sho. \W' 0. i 110. tIl h.' f i*-'et 1 W 'r- xv.ori :lIi.o 1 fo I irt
)W'' oIf C 1t Tr j\l!. N I. CT ri I i .; Arc p resenrt on the lovEr ii tl )1 -s.
Pt. l ps the lloi st i-, iifici it -h;i..ict: 'sttS' o. thIr lowcr rn il le prest c 1f a o I I ori c11, 1 -1 0 Q -n I t li-' 1- v oCoil i of \ I.
.li h. it I }I x t:F, I 1 (ot a, fostCh d hi I 1. tu11 occurs hi i
r;4..I.toliIiiti i ( U dt o Iicr fLiii1 ldlits Lt li c\'c is found ii. ed ti/S .
In. thi. r,. rl. N i.>i Prtl q/ t thct ') .. rc S ( 1-A iihles that if Florida.
ir fu u. eVI< 101 is often i- n ,1 -1 :S o e 'iL lk L il I It is fili t in 1 .t l 'i
fin it n11 tihe J pit r I between fi prot oconiid .ail hypcnid land tihe lower crov.
Iif lit of tie latter. Ai .ot1 her c)fi] l pora.rv gt'tis. :k thokfo:--mI .-. hti .s
: i ,i ili.~ r MNI, 1, [L!t 1n thi. s T i stinlit"., t il-' ftrnll .Ltion o the doublee e ;1iii..I
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vIl, t TI r l ; t! !









7 7
; r ... '' .r .: 74
,*r .
~ f I.. , , . : v , -


7t* 1177 I :F .



1'T


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PA ~ ~ : .7 77 A 4* .
7 7 7 7 7: t



i.) r!IE irti i 77


7*77l7 7~. i 7.

It 1 77:7
'' I..





7~~~~ 4 7 7


i, i '[
'I Ii




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l 1


.7 -4 t 777 7

77 .7

p ~A P:, r :



CF .:dFi d .t'1 hia I i ;f



-I 1174: -

B 1 4
J. 7 177 Ii 74

* 7 7 ;* 77 i;"


IiI




It 1 7 .


, irk .-ii' ni'I' '

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7i 71 1 .













.. .

1 . r 777 \ y, ',\ ,





7 p 1- tr.'r I \t '. u
1 .: .t } T-, . ..





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7 7 ; 7 .7 :7
p i r s .- l


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1 ii Ct.
71 717


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1. i
i i 1
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106r PATTCON: TEXAS AIRTIODACTY'IS 185


ilv derivc d fro t SC.L i'll ti i .1 (',iit Lml1 JUS linlf ii it is often (l etiral c
toi do vs It h.-s hi'teii po.ssilI tI l rt cog -,i, it-, certain shifts iII thke
p;ittP rns of I 'ritliO i('t <'fli 'c ('tia utc rs fi rTuii the (.ar i-i Gui]yv F atina
to the Butrkevili. FLuTia. anid lTi.tEtis( tlhese- shifts ay be irn portant
i, int erpretilng the ,x 1volutioll of ti is group ,f' artiocdactvls, _ithey m.v..
.'ent- iKliv wa rrI ant sI.p rHTa tax on.111: staiotis for sthe wo frr ni t,..
'"ih' B iurke-ville specinmens mar-i represent a tteniiiporal or si1 1.'c',Issijii;l
















3-4
^ : .








\ '
1l. r
















2cm



















F-
I- ~




-c















Sr..'..E 27 I. A t i, RS C n U.-" 1" T' "EC; 31 l'-J(3] :. 1 partial skull and
p!.stri:-b t.il IhV orns.















:r r f, I
i I



T' "'

i : ~ : \

i ; I ';

it' 1
'


, 1 c !


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i i *. * *
4t 4




: ,' . .: --; *',-. ,* < : .' .. ., ',.- 7 ; : :. .

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1-~


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I


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'!! 4 4 4*


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4 12


i ' i





4 A 4

41194 ,i


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FA L1TY. lir.r (- '> Sp'rir.k'u SI.-.i

. A .ia .. ..... ','i[ : '; I 12 ..




} ; .] ', l. .


......... : '- .




Si. :- -I '


.4,


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4 3
.21 4r ;


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. 4' i 9 4I.... 11 ; 4, i l.

.1i1!: IIu11r


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94 4 .


SA ..:: : -. ': .' . : 1..' ,. i. : !, ,

* :, ~;. ",,: i >\' ,' il .! ,'1 : L' ,l l '.i
I tI ii : 1 1 'j" i:I \ i i\ I .j 1 1 1i\ s ,14 1.11




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.. V 4 C: ;


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4,4" 4 1. ,

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'


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4.


94' ~ ~ -.- I 9


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rw l~'i. irt'rl 944.i


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,j !m ,l t (. .O l l l ..' . ] .i'5-r!it i -'' ^' "n. i ;, ll ii -',



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I'AT OVI N. TNL;XA'.S A.HT )1.t AC. l \ l..S


hy-psodon than forins ] ci.: .-r in thei section. Thi. gr. atr' -st c.hian1 gc's
h,-JV r alfec i- the arC'rc'ssor:r cu1sps: the I)pos'Trior .ti i];.i art. no Eion.tg'r
vi.abl.e ad the-' interior ones anre considerably red'iced. The nmdian
piLL r.s, which air 'sp.-c'iai l. prominent in the molars of P. -tcelxai.,
have inide rgont Fr 'iiet .1. e SC7'Ul d ai id fourth I preri'o rs of P.
frt.I'f i.:{ i are tpr .'rtionLa .C-I .siialler thail t OsI of the (C rvin C(;,illy
ari Bur-kt-xille forms.. This is equally true nof the third premolar of
the type of P.. francrii. OI vtL V; r, in uthlcr specimnilis coLifidt _itli
assignr2ird to P. francisi ind :also in the1 l;ar Synthrfceras frlc.frnt.s,
the- P-V i.s iomlonliU.,si v robust,








-. .. ,.^-*--- ".-..,.'.,, .- o .-.. *
l -- r'I



2 cm --- "--




Fi-e at-her w.I preserve lower iaws ( JTBG 321 .9-21- '2
.1219-214, 31 1547. 312-13 :31219.219- kiInon frAin the Cold Spring
Atuiina provide an ade(juate basis' for spe-cific identification. In four
OI (i 114 fivc specimens h1 corl p II dICt'k JTi ti tuI 1is prcsii OF
Midicated, and .'Uist-ts. ofi thr-. pr-i ilars and three iToars. Th-e
($r;iririi fOr PV and d u t hist of thf.S i 1i~.is.riforiT c nine ar: preserved
il .LTBIE 31219-'21 1, a left mandible, P is seldom present. On
1i1TBEC(; 31219-213, also a c'ft iaindible. most of the diasterma and
- 'IIpIys s Iti1,li-s intact T.!. i --ial di; t iatmu of P. ranci.sii
IS C'(MBidOfrll 'i.. igc r han thbJ t If 1 1 7r ., c-1rt .'foIdi iPt r (i:, a
lillr liCif.t -A.d ir d .ia.stenk.t 1 iZi v if i li upper \4 .. Bpc;' ise the
h.-pO sp CUiCrI o01 P. tfxan Ti br kiisn B anterior to P.. thie Itiigth of
Elie ,CI.'sh'Tira ( 'll- Cij t bf:'- dett-r rirlned p )r 1s iv. t, nt j l i li i from sv. m -
MCit F L LO1if I, I I l;ppe14arsc ..t-i i i v sh r r Lht1v hter 1I b a ta r- r forr S.
TnI (li- diastrIjt of' Fri ti .. cra as a4 S. W J.H S ierW -C' ti ihid1 iAorL'tr (hanl
L.tll Of the(.. .sicl-l'c di i font lns. Fac-h ;ra Ti il 0 P. fran li, CxCr.p t thf
ivp describe liy Fr ck (.1937, has a 1. or r an alvr-oh s for FP.


1, 6 I


I ';






BL.SIS.ETIN FI.O1RIDA ,-TATE MiCSLUMN


hut this tooth, when present, is highlily' ri id'd. In 'TBE:C 31219-21.3
are two s I all veo.li for P.. wh.eres UT I'E( 312 1 9-21.4 h s only r"
al] coI Ius. nl i all othLr r.)spec'ts t I. specimens ar 1- v-irtually l Ti( N tic' a
a ri affor .I no rc asor fc. taxonomte sc ep ar.r i inx I vin" cof the
.graidu-l rr.d, ctini of the P, in earlier protoc'eraiti.- i aid its con-iplcte
loss in S!ytbhctotira.s tricoratu. (.i based i1 `AM N 2I46S f .rn tdhe
ClIarcndo(nr beds ., va ri ;iion in occulrr: .'nce of tI is t oloth I ad Pp ,
an4ion, (11iii0 H vid als \,Vit ni 1 i,- v 3. ( p ul,1;Ltion is rInot surprisiiL. illc'I.I
is to hb.e expect ed. P.-I of P. fran ci.i is virtually indi.tiguis ai -' froin
the earlier forms. 'With the., .xceptiont of th e .lr;Liti-. rciirvini L.!f thE
proto'o il., which For iis diHtict r-ictss ii.t i[Ui ll. .-- b t\w'ee -e it and
tI h me-lta c.r o ii:, ,oi .sigk nitic;'ari t i C i ..i-r IL nc in rnor Ip ol IgTpV is 1o: ted in
14. M has a small a-c.c'essorr tu. bet'rclf at the- base if the pustcrijor
lirt of the protoconiud. Tlh Nl:: P fran ii.-i is hii.:g--.ron3Wind :and
Licks 'anl trace of. (i :c' tss )ry' ci.Sp>.


A


B!


cm...
Icm


F iu:.-: 2 0. S,y .th'eto 'cra' tri, rrnatu\ U'TBEG 3113,2-.f'3', rig]t .. nCl.i.Xai


\'...1. 4


I Qo,











S1 1 s "


,, ~ T i' II >:i


| .. i ---: i ._. i :i


1

S' I "9" .i 1 "",. it'

. .. .1 .1 i 'J .i .11 !





,. 't i ,,,i.il~i.,ir Sv !ii t:'r!U l '( t'! l( ;l.lcl. [l'W i(k i i'19 7: -,li. 114 H !IV Cl i! i i 1p 'Ir .

', l, t i l " ,' ,, .. n t Ti(' J.' in ,, .i l,,.i ; ii. ,' \ L ; i Q' -' ." 9 1 I u .1iu.z (' 1 "ti 19. .:
lIN 'l, *\ ]l! '-W ]i


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I ' .- I ", h'" ".


s- '**'' t *r \ : ij .! \ '! ^
: ,, '. I-

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4. 4 .i. . ., :1.


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1: L4 I.4 -i


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4,4 ,( III.

I I t14:


k :,t I I !


il ti' [i ., p tii l ".*-.,-."r L <,. (III.r i I .: ., ',, irtIIl 'i 3,1, .. :t i i ,i; '.l



1 .I.% { rlct.',Vrr t'r l t fr nri! tl t '.,( . !- '(;,t F I l'it j l liiI. li < if itrIr fi

.~t 1 i -. . rI t .i. i'. I 'i .i .>! i ti

f ,' i... p,. nrv it i,, l r' ti ',. -..- !,-, ili, .'. 1, 1i ,i i l ,' i ." s
T ", i> L


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4. r

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59


i;
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190


B ULLLETIN' FI )RI..A N TAIT' t U 51. I N


V 1-1


S tincornatus oo
SL^S0!'^11!5
(Lapara Creek)


P froncsi s
(Cold Spring)


LP JAgn us na -**
(Burkevidle)


P lexanus x A
(Gorvin Gully)

20 25 30 35
Lergth of MI mr mm

Fifq iv .:. '30, I ) i ( l.ribuLti. jl i i ]i .is.T srcri.':its (f NM ., 1 nT iLIl i (.C;ilF C1oa t TIv in h. 'Lt1..-
Ct'erTics.


PIiYOICFXY (1 .TFv S Y' l-i-.lOC:h I..11 .NA

The serit-s t prot-ceratidsc collected fromi siic-'c-siv bioslra i
griaph (i 1 1 Units iln thei T(.xas (.OLastd i lii Ni'i t. i.i (''l.lri U'i r .,s
that c_'cur inT a stcpli.kc proir<.,-siou] tU ou glt to rupris.st-it tlhiru actu i 1I
I- i Opi ll .'

Iivnarkabhic iniforrmiitv 11 this vrnup exists wirthi]11 ach lLaunr,
iAt IJough .S;.aiiple size is smai], the. ithgre o4 variablEitv fo 1t iiiiuSn
spec' rimis iro I withly csparated l Caliti. es I U; is c paral'-e to that
N.pu ts('it(_ il *a sinlgi m- d-rn popsi]aJtinm of rii- r ninatt artiLii 'dactvyls.
N ) extrrPlite V Arii, ts, iA-\' ht.'cii d-rt, U. *f'r to iT
  • Of divercj'"it u2)gL L IiI V .pit.ratfed popu la 'i ,ns. Th.4.s it appr-arsi
    ti:ait theSe anitm als. at a[n nl, tCi 0c \''-."('c ititi i.' i,' li_-'s ol ; of la f t-
    interri-et-dirig populationn. or periia.ps wtr dcistrihuted ii v'ery closely
    related allopatric pop.abti,-II is Iwits oIa'.isiorial oI epp-Fort1nity or con tact,
    prevent aing th. ,ri i i o.,n, u1;ati j..f i 1 ,- lit- im.-tcchanis1ms. All evidt-'nc

    Burk(.'ville, (. l I Spri ng. : I I ,.para (iCre'k FPauntas represents .i
    C'_Yl itii ,lVlIsl _V oIV( I l Vli t J, I'ii ,.A,.U.c,_ S. .. .... : t('C Si'' V it' l it i, l ilsOt ,ilt( \l t j )-
    I.ll;tiiorls. Xw ith _vo)v Iioi J.)rt c <.'i' is;. i .'z IN I v .. ic-t. :istainIed iuiri r'it
    te. ro if it .' ) i n' l\ Io s it o(f th

    adaptive optit[nri n.

    D ispersal, w ith iIlit, (, vi u f:.xc ci. ti ,or ti t i i l ;i[ IH i iilitii
    into hi.' Coa.stai] ilai t-)ipparently .t.iis, on, \ c'i.rtiioi fronm thi:i.:






    10 PATTON TEXAS ARTIODAMCTLS 191

    region, as no discontinuites asriable to replacement by adaptively
    superior alloehthons are evident.
    The earliest protoeratids are known from the Geat Plains of
    North America. The genus Protoceas is generally thought to be the
    ancestral goup from which the later forms were derived. Among
    these, Protoera asutus of the White River appears to be the most
    likely progenitor of the early Miocene Syndyoceras The skull of
    the genu Protoceras is characterized by the development of bony
    protberances rising from the temporal crests of the parietals, from
    the supraorbital borders of the frontals, and from the upper margins
    of the maxillaries. Thee processes are rprented in each Protocera
    species, although suppressed to a greater extent in what are believed
    to be females. In P. cele the maxillary protuberances are broad and
    platelike, acve outward, and are widely separated at the base. In
    P. natus they ar much narrower, more rodlike, and umte suturally
    over the naal canal (Scott, 1940: 577). It is in this character
    especially that P ansutus appears to antincpate Syndyocera The
    gens Syndyocera has four prominent horns: the rostral or maxillary
    pair join over the nasal canal to form a common trunk, then bifurate
    just above the juncture into two outwardly curving branches; the
    two frontal horns rise from a point above and behind the orbits and
    urve gently towards the ndline. The kind of postal horn seen
    in Syndyocera appears to have evolved from the union of the two
    maxillary prouberances of the type seen in Protoceras Msuts. The
    frontal horns, on the other hand, were probably derived from the
    union of the bony processes on the supraorbital borders of the
    frontals and those on the temporal crests of the panetals, forming
    a single beam. P. nautu states those requirements necessary for
    being placed in a morphologically ancestral position to Syndyoce, s
    and the later synthetoerines.
    The lack of a sgnficant number of frontal (postorbital) and
    nasal (rostral) horns in any of the available collectons makes difcult
    the detection of any evolutionary trends in these parts. It is difficult to
    determine the constancy of anatomical detail observed in ontem-
    poraneou, specimens because of the very small samples available
    from any one fauna, it is especially difficult when comparing speci-
    mens collected from each of the successive faunas. It becomes easy
    to cofuse variability with phylogeny. Notwithstanding this m-
    plication, theancestral horn condition for this lineage seems dis-
    ernmble in Sydyoceras The postorbital hor of Syndyera extend
    laterally at nedy right angles to the long axis of the skull and curve








    up and towards the midline in a wide semicircle. At the base, these
    horns tend to be shghtly Hlared and flattened, but become more
    rounded toward the middle No bony protuberances occur along
    their edges. The only recovered postorbital horns of P. teranus share
    several characters of Syndyoceras. The horns are directed laterally
    from the skull and recurve rather sharply toward each other. They
    differ from those of Syndyoceras in the slightly greater flare near the
    base and in the increase in irregularity or roughness of the anterior
    and posterior edges at the flare. P. francis shows considerable change
    from the preceding forms, the horns sweep out from the skull at a
    low angle and in a posterior direction, have greater flattening towards
    the base, and the anterior and posterior edges are marked by more
    prominent irregularity. This progression in characters of the horn
    continues into Synthestoeras.
    The evolution of the rostral horn of the synthetocerines has in-
    volved primarily the lengthening of the main shaft, resulting in an
    increase of the distance from the base of the rostral horn to the
    point of bifurcation. There also seems to have been a gradual increase
    in backward growth of the distal, or forked, end of the rostral horn.
    This is perhaps a counterbalancing adaption serving partially to
    overcome the disadvantage of havig such a heavy weight towards
    the tip of the snout,
    Thus from the preceding discussion and from the data included
    in the systematic description of these forms, a morphologie, and
    presumably phylogenetic, series representing the steps in protoceratid
    evolution from late Oligocene through early Pliocene can be estab-
    lished. On the generic level this series is as follows:
    ProtocerasSyndyoceras-Prosynthetocems-Synthetoceras.1
    Several morphologic trends from Syndyoceras of the early Miocene
    to Synthetoceras of the early Pliocene can be discerned
    (1)" increase in size
    (2) reduction of premolars
    (3) reduction of intercolumnar tubercles and cingula in molars
    (4) increase in crown height
    (5) increase in diastema length

    'As this paper was going nto press, Stirton's (1967) desrption of a new genus,
    Labdoceas s published This genus from the middle Hemngfordian Flint
    Hill fauna of Sth Dakota, is in my opinion very difficult to separate from
    Prosynthi et .. have not had the opportnuty to study Stiton's specimens,
    however, the w taxon is disuossed in Patton and Taylor (MS)







    PATTi'( )N. TEXAS .\X !I (! ).A(:Ci'1 ,S


    1i93


    .: 6 j mi i psti'rirlv directed post-rmIial iur
    i"7 ii increase it iirr egilaritv of ]itcr, edge of


    iS inLCrA.t. ii (d istal rlo' [on 1 ) se of ro.t ra


    ,!1( ) r l'd i, d Ve\,]oiptT 1 ,., b(.,nlg v (.1 o; 'ri tL ,.(
    n(id trl0(od viM e .l _upp.piBnu 1 istral huorIs


    postrl:ita h-orns


    ilortI to point of


    C I1 i or ]ieIvS


    Si frat order P oE <-:() KA L.i -nan ,s, 1 75.5
    Family Ci: vipru: Grav. 1.521
    Subfamily PA AFO'. -- N Ir~ l Matthew, 1904
    1as.vtonu rux (.ope. 1 8,77
    Ilausthnrrix rle-ans M athlw a: nd Co.ok, 1909
    TaIle 21.

    Tvp. AMNIN 1401, a partial left rmus, v-itEh IP:-M,.,
    Oc::t.ccLR:E. -- Lipipara (Cre:k Fauna.
    RE-FEnnF. S .\I F -.t UTFBEC :31 132-3'"-. pArtial 1ft unus \t I
    M\'I-M .11:32-4, d) :30596 326.. prli.al right rain, 1, vith P.-M.-Nf. d
    broken MI.; 30N96 496. parli'l riih rinmus w ith Pi-\1i.
    )FCt m-' U -Io.N Thlv -1o [I.r rt- .r,, ..;p(D. -'ileri nlls fron Be,---e C(i-lntv,
    T'_'xas, are- indis in- gl.islha al f romn a series ( AMNI- 17345, 17341.
    17.'47. !7.'48. 17:9 I. collefctd from the Snalk Creek beds of Sioux
    (.C ntiNitv., Nebraska. and de si -nId h\ Mlattf ew d Cook (1909. a
    Blastint'ryx rle-.ns Frik ? s,-paraitcs tlbe gen,'lra BlHsme'r
    ,Li. L oniro. rume'ry. on t ice basis of the latter's elongate diasteina
    a"ind Tl ) rI. 'd. dP, r l.-h-Pj. No syrnphy.sis i-, preserve. d in a Y' I<:f tie
    Iiapart Cr.c' _. s(pcciin-iisS, .uit the pri'u- -ILATr.N :k i- t m r..i k d ly rcduc'd
    LS il .sp, cii .s of L.t'.n:i r.s.tromcf trx. P-i is r-cprc.-.scnteld b. two ai lv oli


    .ih',- i 21


    \Pe. r e eI r


    ngth -.l~N 1I
    it .It'tli M i-M I
    I:,. k-.'ithlt], '.w fi b
    P[, Ivrui wid{h
    I:. I ITi" 3' 1 1 R I:
    -N th *rjj -
    M:t, lt ri tl;t a' .hjti
    M iI'L'rb li
    AF:r.fl liliihtit;-totJ ( MaNinlt .'.w'
    Ar'r:oxn.natC


    ..\M Ni 3 11 E .'THK.(
    r j"i :. i' ::',[80, :l C%
    .:NI, 1I ..1

    45 (P -1
    '.-.U 30 7
    6.2 30 65 .- .1
    7 .. :
    i H) .. '2 S. 4 7
    i .'. .') I:J li .. 5 .'-
    1 'r"l J c i .1 0 ..--
    anr <'. k. "' !i g. '' ,


    1.T s. 0.6 .2
    30Q.S-3fi-:


    L 1: -3B


    .1:3 8


    S.4


    . ,i".
    ", ,


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    5-


    I2A5


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    D~~I *


    - j. ~S


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    1'

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    cm


    j.


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    1.

    i i : '~ i i i i

    4
    1::

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    i
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    i







    I A T PI IL \ A. NS -\k!I(' I' .) 1A( .'lI Sl


    S T h.{ l ", [ .I' ifrin" .f \' ;i.,, nLL d 1 Frick li'- 1 i

    J.. ?u'rr.iim ? [ 1ck, l t ; )ci. f l' I :T"i ,r Trx: ;i'* ri ., /.. ," rir) J sv
    F-ick: L. L Ji r e r l r' l ,t r' ris )i s- v rT, rick, (. vi, r-
    eilt I Ha; d ; /.. Li rick. I c n ie i no I- I itanti i d,'-
    -ii.e 1, .1 ,\ I i I. nt i n .. L. n eTrria i i7 ;ii' i, idr iOirIp!i O V i ind
    bit ivI t it i t'Il i d 'i s .siO i 1o /n il, :i c L. 'c! i.
    1I i c u'tli (t IJ r'rti!..r.'t' F Vt- \\
    ;.-'1 ( Ili I .Vh i S <.1 IriA. 1 f iei'f i/!i IE.l 1i I -.li l)c i II' IIi l t (itn1 i vHiit
    ' l C Hp.1 a t1 1o.1 ; / .11d)P
    \N t li iJI. Species
    .. l t.-i'~J1?i':.rtU ";.H' lS' r i'_111 til.' ...l h lt (_C[' 7 4,'eJ JI N N M ixs. l
    hi, .etl a : i p l.urti ll ft arnus with I P., alv di a d I4 1 C i
    -titlrf',' Cit ti.- t\\ f- (liAgnO.,d.ic <' _irLctftr.I, of1 Lcm. 2.iro:.trun! .--r!r., re]dii
    tI1Fo .:i 141 ..ti l i 11 n 1 1 ( t'N I i i l pC) I Cc (
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    IPAT [' TEXA, A hT ,I( I 1'A(_.TYLS


    1 2r. Y. III xr I (1.rr I t Ps I Ir
    :is D ra f -r/. Ia. E ;.htic EF c'.. i 1 37i lists as (. r,.1i'ocC'r..' atm(.v f~.i1
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    r(.,tU. ,ih c1 r hi irmJi. t is Fr( 1. 1937
    I'iuiirc :32;1
    [ 1 T-r(. T
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    li FE-;i -..'i). .AT IU-1L. ITTBF.C ] ,' 1SSl-14,O. tz i,,u n iattr, du..n-
    titin, a; nd 1 ; TBI C 3 11 2-455i. ipartiail horn.
    Ocu-inp ?-I r Lppra C,_re-ek FLIauna.
    INrX f i'n ioN. An Iin .ature :lntiti'n 'TBE( .310S1.14s0 I
    rill B 'o') o .i iv, Ti..iy, .s, .(rc..T-s \v(r\ tc ..1\ iII si/ tIrd(.l u p TIt.)r-
    phio w itl a siia d.tlit.i FAM I:32216A .. frovin the M\ueAtdains
    (itiarry, (l;LTrefndn!r. Tc( is. Oth i.r n nAitiAtii're Li-'ihtti n Ofls of similar si/.7
    .. FAM .1' 2491, Su 'Ilr-murmu .rr. ..roti, t dii FA N I '31199, Bt1r- i nI wrr,,x
    wlrust',"'i.i ifl .er itU..m the L,.ip'Jir; .a1d (C',rxndon spacim np pri-
    Iriaril v in sizt- O i ti rrcleR 1f 0II; drin c in Ihe pr cst- n ,f d: "Pa lOO/'f-
    merjXv fold" on M,.
    The left postorbital h)orni, UTBEC .311.32 453. from this fauna. is
    broken distally. but on0 the basis ti_ l prxii ial crox, S-sec ti al sh.. -.p m
    1m1_d t i1 i" r(ei iri'f f rl ; it](l ,tp -r t, I l, irT.'sr' l p);Lt o th slalt, it
    mnore o.slv ppI)roachsI tl posttL'bital h ors. of ('rioC''i'i than li
    fromi mi'r.x, Bio-.rornucr. or nv other similar forn. For this reason


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    PATTU..N: TEXAS ARTIODACTIYLS


    tripinnted cu'rotch.'


    theli sm ill siz e, tC e id F~l y p ai i ni ion 1u wd ii j.t it r e r i .Ilr poi ti ig.
    fliev i c alt1, ir HiiOT nTR J.i .ir\., iniiiatur, l 'oos1 zntil'rs..

    Nlervcodorlit i1n(t.
    Occu -:n.Nc:. Lapara Creek Fauna.
    Rr-rF.~vn i M ,,FiHA .. .:TBEC .31170-2, a riglt \1., and IUTBI(
    31170-S~~5 l. t Il,.
    lI.)sciulprioN Two sminali hypsod:ont M.' from Bee C(A.nt,
    Te',Ls. itre II rred r- f th nI Ifrv c- iIni oae. t 'lial N ) gini nric d(esit l .'i
    th.1o of these isolated molrs is virtually iTn possible lnd is n1;t a t-
    tempted. Whetl.r tlI'vy are' refTralrh[ to reco ized merycodlonts i-
    (the Lapara- Creek Faiiiit, e.g., Ramocrrros. or represent .an othenvise
    Lmre-.CCO izd group is equally uncertain.

    Infraorder O()-:vopor,. Osborn, 1910

    Although, the prtesIi '" of owin.' .hTIt nt'eri; L in tlie Texas Coast.al
    Pliii lauli wa.s prtvitmsl- reported Iv (.)ti ir 19155 w ;ind lby Schultz
    Li', I1keibach ( 1941 a the< frTiirr di.strtiltion of these animals
    tin t.e Ce I f Coas?-t theiylli JLt I ir llllicIi Irde2l'v Inknown. In addition
    t tho i thrc-l Cg IT'a T n i l' i \,.is d.t' i'fl m'tr iij I t is p pu e r, rn-odonts
    nr,, a-re kno-wn t: ci 'pr is lorid- r!<"posis ;is we']l Maglio ( 1.966)
    dc-.rribes -!.-rr-oh/(q- 4s, tfli Thoi,.t,, Firn,, wh il,. Pat-ton ( 1T967,
    [966[67] 3 r]pi,'Orts -r.,:' .nits frc ,ii ,."v.r.w l diHFferent l'Iorida localities
    r:tljigiri iiiE alr ffr4m Ia. 'i Olidiosen$ i to Middle Mio'if(nTl. Aside {rom
    t .i' 1ho.'da intport.a n(ce ., tI S ct(Ul it l C',s ci CllloQnIrstratt a r g -ats i ac-
    n si 1iitV ofl th.1 (a. l" (.o. steal PI .tiii (to0 in I rnir tion froit il, ( .reaI
    lIati .s thanP pr'vio'i lv ha.d be.'n COsidCertd probtIle.


    -a mYi M I fY(OiI DO)TI11 ,\ T ho rpe, 1923
    Su. olV FTICHIOLEI'TINAE ScIi.t/i. aid lFakenr ach, 19411a
    Tichlepi. C.oope, 1S7S
    Ti.hol.cptus ri'e i S-' lS z hu .l Faiktlk en hac hi, 1 941 1a

    Table 23
    TYI'v: FANM 42329, a It-t MIn:-Ldil(I) with i1 ( al.)-M. nd partial
    fTI' U1 .


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    PATTON TIEXA A. IIOLLACT'I .S


    MI-rh lJm1 ininF1. PctrIson fromiI the 1ow\r MNI'arsland '. SchultY
    ailnd F'a]ketri:.1iach, 1947t' .


    ACE AND (CoiU-L..ATI.'LON
    This paper, as one of a series of taxonoi0 c a.i stra.tigraphi'
    studiess of the los.si vertebrate fmri.lis ofI TrhL Iar a.Ae in the Texas
    'ulf Coastal Plain LVay, 1924, Ilesse. 194.3, Quinn. 1952. 195. \\'i-
    on. 1956, 1957. 1959, 1960), provides :tdditin'l irnfori:tion usenl]
    ri r ;akin3 I,.. is Itr,:tti rap;phic'. i 4orrf laticn r bn I tw\ .en the TcxIs fa m1.11;s
    and those l-stewhere. As er]h i.Jtjor rtaxo oi-ic study is 'oiplt' eCd,
    and the systeIrn:tic rI-vla1tiosiips iof the various ro,:ips t-Iecoirie b,.'tter
    k, foxx r, i'iH"rc.tsi i i[!.f'o Unsi.-itii, i which 11N .) base studies of .hiostr-tti-
    graph-y, zoogeln rply, 1111nd pal,-,JJec-oic.,,ry is rnadc avat.tbl:. Because
    t l(. falin ls5 ( 1_'1.sCI.5 XV,'i.lso 1.959 iri t 'esti.on correspond to ,what
    ITiltl\v )iOStratigr.:ipI'.,er c' i ri ass ,en-s ,i a--h Oi 1nev inlh I tA- io%
    pe viously poorly known coInp.onents SCrv.es to b IroIadenI tle founda-
    iI l ti poli w ni,'1 t Ile a'~r i t areii c LorL.' r-.ists ar -d all'eviatel ovLe lcc' -
    pcndtcice if- 1 r : i if-w tit I r (Aorrclatiin piurp loss. -;vIdi-(:. pr'sinted
    it 1 thi is ,stiddv iidi(';tt Coastal Plain 'ai fanac.r b .. modified ( F.g. 34 .1

    CAVE\, (;-1.LLY FAUNA
    Stud 's I)v Quinn ( 1955 ; and Wilsm i(1956, 1957. 1960. estan
    i.shfd th, fir.t ceaUr outline of tthe' gcnI.-rA:l stratigraphic fran we%'work
    within which these faut.nas occtir. ()uinn (195.5) and W\ilsonI ( 19601
    (o1(.idered tOW- Ga rviln Gul-lV F.uniia to bt of late Arikareean aLe. This
    view was based primarily ,on the c'losW sim'-ilatity of the c(arvin1 Gfly
    tax to those from the Florida iThomas I[Far dthposit.. The Thomas
    i-ar t at t1haf ti W;s Ihlt tlo(1 iin i hi Iit.W llTorne For4 a tit.1 ot
    ri(ddlei Miocene- ag,-. Apparently I,("caJus' SiUipson I:12, tlAi.vxed
    ma,"tria1 froT, i 11~ t. iwltorn; Fori-,ation., at Qtiney arId MIidway,
    Fl. ridL to surnewh ,I v ni r thiLan (ha i t fir n tiH. h 110as r,..Ti.
    u1.1-.rrv, 1.11ui n su., .'s d t a tha tl Si. T 'ol irr .s r.inn fosiI \W e prol"-

    ['hr \17iarsi.mu F .'ri n,.ioT'n '4- SJ'ti,,lt I ,.- 0 1k lk 1,b.) L F: i l'LJ.7 : ri"lI ti ,t-', ii .i t llia
    Pet.rr T 1 0 ; -* r i s. ," .is t-.r n: t i. )n -rprr-'Tccl )v M ..;,k 1 6 ,
    19b.5 .N McKcnna : 1I? l, .;: l SkVinn':r i ners. e rn . Tn "'Ii vrr Miars.1ii 4 , Sc j "]iti 1i-!I, ,lk nlih,.) h t i. l. : .ii. t, P E',.-rPr 'r I p r .irri.- ,rii aid th,
    rr trite t.'.l \fM a- r ,i.l ,d -..k 1 i .m i. \ I. KP'),. -, l" .* '. T I. p "Lp r Nr NM r<-
    ti:,'i i J1'Ji4 gig -C .(, I.', lniriiir :v_. ;iter F ',)rT i.,tl.'I1 ', S .e (K'RA E tL, l-jf,5. r it. '1iCv'
    ot t li pr tl rn. .I


    3 WO


    20.5







    V .i. I


    rb I -.L ['II"N I L, L..)Ll )A '"TATE li."S U I


    a.I v 1) -1n111Pi or carIV M\ion-T.' iiter. The ins of" hort h c I Ltiw-
    i}IO ih a dlli Tinaf.t If. i I triti L-. t t' i .l-'r ill, t i tii ,iis of

    l 'r 1n fll 6.i . iil I- th i .' ( f. ,' ,i,.l.. t .\r'. il i _itn". 1 i tiltI
    tIl Ili ii ]h'. 4tr (-I. T ; ii s eIrlI. rol a t
    lieud initoi rthe terrr's1 i;).1 N erttbrjtt" idLi l. ThtUS. II1ri' is 20 \'i.iCIT .
    it'r t ill iit it i t jii (ii'i f, i (_ i Slll ti t ii l It 1 l i it iT
    tifi t.l -ll 1 T i l 4 t' r 11i r r e t' 1 0 1 lif 11-- )J. 6 'l"t
    t10 11 iI T11 t 1i fllr. III (14'Iud I ri' teric-hrofiT v I 'K
    e til. een (I Tl it'u.c
    I lls ll.'dti sl 011 ar d itp ( r1. Qtir!ili t 1 ")i; 72 76 5 ,: i ti

    , -t,.' l hu i (.: 'l\'('CT Ni 'h" '. Tli- Tl > Iiut., I- 'ri d<,-pn."sit der iitl-. v is
    11ot c orr'.tiv \ e i t',.]hi p.i l-z s n 1 j i l. s ii -C; :i it I Lcd
    d~.-1 I L t U, ;.' Iltt ll3t cti ill -13 l; t Iivt i ar.,. i<. ilt i..i. i ff Curl
    11tn il C. < '-) t I't i L _i"' ait)ti iE! \ i r ,I i-i 1 t .' Cs; tt c -l4 1vt .
    .011i t witlil 13 n !,"i <_I at 'hI ilijl', F;irili. i l' _' l'orii- ( i, on]I jic'pj l
    S.1_ v iI-lcllude, h_. Thi.-vnIL-s FaRtaN sf.'!ir-n nts in ihe "Al.._h. t ', Iori
    mn tti i 1 t' Cl m 4i St t c( i1 i ti f(I

    l T' i"r iK ,i l,'. ur', ...rlr : iI .I ll l.-ini 'h1,,' ii1 v ." l, .I
    .-I : I i r'rp ,.- ,L -. I.. g.-' j<. *-' . ; '. ., , 4,.
    -,,I t T H

    LI:., i~l tii_ (t" ~~ Ii14 ." 'NW 'tII 4'i l l I' 'A'I u ,r, i' lr'. ;


    .i.i,:1 p i l l-.- t. ,-rtr I,--:I r lrl iir-i' ",ic ",i.. t r i-;J l'-, ii.. t. .'u : f,.t'L i ii ;, i ''L ,Lii ,,
    'jJ ; i" L j'i'< t';' 0_'1 tr 1lii j suI r- s .. 1e t ri T,,-
    j I ii .1 t1i T ],.1 .








    tflit
    L 11 t it .Si i L-,. '. ii- r]- (. d'' ).; C i- i I-l t t ", '! -lI'tl J"1 i r ilr ." '1


    O t ri id t I LI .'t r; i .' tI 'tlv It' i E 'i 0 il0 f1 ItI 01 1u ~ ii 11 illr ( li t ) 1' ,; .

    s iL sC 11- d t li n- I n thit- -lei i llit Ini rn. 1r6 4. In Fn t it I.
    (.) iii T I tI Il Att PrI ,g il -i. %IrI'L rl Ii firi r' sip tSo T n !' ;v the
    i V toISrW f A11iri otl ISd 1 t hIlt toa rli In aci.'c r111ud ittd 111 (I 1 O[t[tIfl
    hkilid of kaTirSt-c 'jntrull Ld :.C;.ttC i \i4' ati b;s t siit Le iO tis pr1v ; c es' S b:A
    is:,, n ) ,.dlp (i't sir lc. -1i l .ia O: It i l',i *tf." dl_..tvsi.l .i er,. 1 (.itoi

    t t il 16 1(', I I_ t t ',. Hj' ll[.l.t 1-1. ,.ll( ltl..i ( i! lI\ Ci 1 1 14. 11 il v

    ,c-il-t~ptiLd e of ther te rn (.'l i'. Strat. Nr' n .'nr, 1 61 IIn. a veriiit.
    l'('<;insht.' oitf th, I p< 'l.(jT ,O 0. 0 Po tt hi TKr I l ;, F;t 1r
    ,.0"lT- t.r .ri.,1 i',, E t- t.t"linp L i;._ O .irhf-tv ii < 'itOt-,ll" ltii- ( .lr

    T tc ag. f' o..t I "] Tho I I I ri- S I;.I- 1 fnn1 a g(. I i -


    IST h I< th p c It f 4 -'1 1it .n li.i-. < ` r.i! n' ., n- I iI U".. 1 < irt lk e. t ., lu. i rt-n'r.l
    F lri .I iI. YT) `, 67, W )


    2'iF







    1.9i9 PATTON: Tf .\'XAs, ARTC!i .DAC.\ I S 207

    sideret.l to Ib AIt rikircur,,' 11 -hit. 1[4 -' l i. r i t t '.. hr w -14-
    \v, ,-I U cr. diu t 1h ceCi hi0s t Ihesis 01) p;L 1 l .j1 'h `.la phi. I .. aI en s
    I .IlCtfItd it h 1 (t1'dl, dep)1 k.I kd ii't.h of 4 I. L ti t 1 l,-
    ,-o L'.' d I()k.'ii['!..,'_' T iu : tlI. t I:n tixt \,. kC-r. ,]iskl0t1'. itp.,. Ir
    I 195 111 1t i;t ti f t Ii T hoit s FoA 1in liirLMs 1 -( ii t1 i ,- t(. n1 1 1
    'is l,.i si, lit \ A .lk..r-'," I,,t II:'-' i(i iii'. iri u iw i ill pa.irt in Q Jinn)11
    SI cst. n o tin. ;L.!' <.ff tt,.. "I 'si;ii'c _, Far',-m in nrel a i -i tAo tl f
    L( I. z -. r.. :11 n I. -k t -t co I l
    .irv! vi Uit i FCatiii ,I .4 ai 0w r rf t clitit p .perl. 11 E. \V.wod [9-4 1
    stut.' that .i ThoT. .: i rm hIj i ,OCrIO," ;n s, rDi'rc, te n ah r. uri, is
    pr ,.' res-, \'s 1 (. t)\ cl assic e1] V N i.r r i ict l'. issi1 fru t th(.-
    F'irrin ,ri r d.-( s. /j). If. i, i ( 1i itiiii ;s :., '1u Ai ik f -.tdil o 2 mair lv
    I It<'iii or!i tIia I I .I i, tE i. i,i-]ios it. Oti the I it s ofd their cIoser
    pr:o\imii2 to .N ud ,r1 1 :1 tcr n111mb.r oi taxa shared witi tI ] Iv. 'ci }ono-

    1 4 iF II ) ('.d1T Isc Iw, t e.l. tzt N
    CO is' W3i 1 ..i ; iapprL-i'cf is I (- plai' ,Witl,'. ir. -Jre i I1 tTe is
    ( ii:s;l a iv n 1 n11 s (C -tin 11 ti :mas lid n, th tin lI


    tthan tT<- I. ..i o
    11 1 57, 193 I .( ;11 S 1.ei i r 1 thY (i t i-\-ii (9l IFaIx ki tom
    ti A r- i k n l'ntssis t f )ah ?i1.4h1L ci. cIS i)iuoiruu
    Itim r Ij.- s la. hfi. : l-d; c'u__iNi- t '] -}.ice \. [i pi'Jt ii-t. out th,- t tOF i
    :ss<.e r 2(i,ui t 1i /"i I/hi c .' -I( P Djruj /ll/Ti !. L ;IT rI P TU /t ir.l1 L t is c'Vt, I'Crit.t -














    .. .
    th1t If N_;ll
    '.. i. l Ar i k.. Ic i n, a,, !t.si ,E:itt-I h-, th-e .V -;d hh( k.' rni i- it-t e \ I' -Wo i
    191t 1A 19.l % i,. 19.59 ) fi rthI'r tsL- '-id tha:t o- f t },, h1or-s' rri:Itci'r Ii
    -i j" r 'Yi pju.s' c -and p !.)pCrti'!)k.s I roIt f i 0 (1.1itvi G ullv IIuIlU,..,
    IU per I-,.it p .ssess M nr1//iqm '. A-Iitcrr.t .1S, ,tI W iicl' iS $I lih['j ts) I':_

    '[('a le, 25. A(;. l'..AN '; i ( .UIVIN ( d A. i'.'i {; ",'i ii',A
    F A ,n .\ rii..:r, .-.t p I t, '!:ii: ..fi ,r,.] i ;,. ,r a C Ia u I,' ,,, I I r



    t 1( us
    lP hip',.-tir \


    C.l(i T ; 1 1 1 s
    / )j*a, 'rj ',j ,r.
    * .'\*L; f(. ,i.v XN t .

    / ):,- r,'.th,.'ri',i u.i \


    H C.,.' hipLiJ .S:. .. x


    .- if .. ,in .
    i t i f',, ,,-(,o







    BULLETIN FI.OBiDA STATE 1\ISE 'i1M


    fii-1(n 1 f (t di..,dr (1956 1 for the Thornmas Farm ho rses. A i on ii-
    dividia ,i I lI s of ti hse' iorst-s frorn the Thom' t. Fartil. several ti i~l
    dnltiions exhl 1ill1 in ioth Pa ralhippu a j id :A Jr/chi/f us tfeati res
    (White, 1.42. Bader, 1956:. \ilson (1959: T77:, foltowtiu White
    ( 1942 and iBdr 1956 concluded that:
    SitU-. fParaJi Pu, ,. Ih ;l .r i t.ti .- I,r1'H rk -viliv 1-':i1., f it i f'IW ,4 'us that it
    l i it f li tt'( if i T'r '\v; .A71( |S\ i i. ,ii TiL lr K>}\ t x .r\ i7 p1i 1E L. i
    It F)pp .iriif iicet- -W e \.. f.- ; .' l I. pi-, CL l.II l i'l:k ii *'r l :
    4 Iill, ff 11 ilt a n f, fiv1 1J m In t t1"a K .1 1O1 UIii In Ne I Flu I p5
    ,.i ,l iL I (>1 l St L*', 1. -Vi i i'l l 1.:. 111.! -.1 11 '. I
    Coninuifll ., si (Ice th- iT ppffl~ i 't C o' I 1 iin't ftin, l-l ( iX fr/h l :, o.
    tha1t stLagc of horse ev liition repr.esene td 'AM rychip ." .e.
    presunci(iC 3- (:rf '1111*Ji t c' i T -t l' crrlc4 t- I. tic.. uLtrks din I )as< 41 t ir
    ,fif .,'i .rt 'f-ordian.. th, H -.tifor)1'(T tiii It K, 13x ,\(tintri i ; V tJra sitis i. .) leadinhit
    t) it mILst lo ia Iy prScedIc il ;Ils retai rdc s .s ,tn Airik rcr-'n evi-t. Thu- tli prese-ce `(. 01 Da p/hf 'nv. on
    a d I )i u dhltu N, flr it I Ir J- f wi -ii'c is k11 n ,i i.t )ve the A rika iree ., iand
    [lit, Far'alt hiij L-A,.'T/.hl;.i~.UiV tirai lSitiosl. with i.,,s attind.-trit cvollitti E.iary-
    zn.ooge--.,.raphica iE ,pl! ti- i s. l-d W i]son t,? pla ce tIi[ G.ar'vin ( uillv
    Fauna in lthe a1te Arikar"n,.n.
    Althoi. ,gh t he fir -st a ppea ran ice of A -. rchcippu.rs has been tsed hy
    -sri 'lKd' 1.i) to00 /t' tle ..is I H.iei itifr._ dian i I NI ri i A11ic02 tri
    It'rfiarv dc. posits, 1n t"vid,7"l'1ce Isio1. this single c'rh i rii to 1-H valid.
    I1 t t liUC' getI Tr;.I' a'cc.epted 2ltiod t. t} ilon ioll tchit 1Li-1 alf.l C Ffr't-l 1011
    of the North AmericaCn iti n'it;al 'i'cti..rv. \\'..od f; :il. ( 1941 i mako
    HO c'ini fr (f41rc4.fippiu s 1.. marking thl b.i.-Sv of i1,, Ilernmingfrdiari
    but rathL-er .s Ji.)1\ list' it.s ih -.t f'r .'cf .i.rrlie.- ;, s i -c iL9 iIn tli Hel'.i.I_.-
    ro rdia u F.u'tJler, fl.'rl hi.ppns doe.s iirt o c-'t iir it lie hI -ts of t he
    I nemin.fordia n in the tiyp a.. t're. but actul] rplacs Para iiJpu-
    weHl lip in the IlheminciI -rd scl frni.i. InI .1in .trl work (o)s .rn I 19!-
    9.S-99)1 in his actionn on the .-1neric c dlian.sis: of cML'cr1"rhi ppus, I isis
    thi:' first occurrence :f t:.it gc.4,S ii the G;-a;t Pla-ins :is b.in i th
    Shep 1 C',:kk. .,Morri.s Skinnm-rl per,. coion,." in his ,study of the
    T'c.tiaryV. hors1-'s 4.f t 'he (Gri-at IPl in.is, also cuntc..'Ids that .AleryclI j u
    is not found in beds older -h.. r SIhp C ( k:. Co.k i. 1960 '; ioUWvcCr
    t'cogniC,1i/ f,.s f-'r 'i t rIy 1 \ in Ith Il. derli .in Ru iring I vatr Formn-ti n.
    iVI.(2.N MNcKnna ( |trs. inn. c.airis that Co( k misidentificd a
    V' )Oii 9Pr chfiartil F 1 ( .t ii t) 111e Rn~ rlfliflf~\\..ttf- f-.S ,IT l H ntolihiiiTwiU1r
    itself. ; In a prelimin r-,- p;ip horse' eVolutipn in1 th; Great Plains. (Cook ( 960: 20.' i states:


    208


    V\'l. 14







    PAT'I" )N. TEXAS APTFI() \(_ TV L.S


    In the- l bl-, rprcs -" tir.4' thh,- e:irlhr st.j es of ,. ..at is c urrr.utiv b--ei'., c(',-icd
    t t'l .uir ird i:'r ; til LIrI t I i. 11 I T rste : t C Si i p1 f!V, I p1frt dS. brC. l'I I'i'.

    111 't II
    Hi,,'Lhrs A it,-i_ .it.'b 1 t1 l J_..'[.i ,;3_il _'-. ,,,_ O .. k e -.il !.

    "f' 'l n.' vt 'w t .- t... I.h ci int 1ith 'r It.. -' l i.. d-Cp si"l,)V.. 1it .. I t i-c,-
    4.' t. .1 ith N i:;j'k"t ( i n :,i. l a.i) 't. ,,d l.w s.6-'1 11n (l 'pt l:. mtd kl, s 1 1 id 1 :1 11 r'.
    a ;f:,*'; WcCt&'ii.-rl' N '.Nr t .1 1, th-,-, ,w. dfcpI juits,. I iv. .- i:.t E i, ic- Irm- ir ppri t li
    l't t iY \IM. ; a.slai",[ i,, iLttS,. ii, "f i '?:l:,St .i.r.:f.. t iir o ,i..l( 4.' ,t '".v',jiih I ii ',:i n' (i
    it ie l, .li, '. rI':' c '. :i f ti' ri- ;,ir r...pi v h "''re mr hi'..' 1it :tr i l.
    ltt in..rn i. .'t. ,, l Il iL 'l' i'm1. i m1 p . -'F tll- .t1> mm r I iN C l i( ( ii .-
    fir-.t tin i ,: C C ;iC.' i tll i Te l ', a]h L:: ti t .('t( i, v- Iti r l iI-If ,.. '4 ";r--il ;L'itji.i1(
    tileT- .:". ,\', v, }: [}tri i:., -s 14 hi.u !;i': r .. ,F. *the 1:_r( 'I: v :-;. \p, ,' -, il-'v ,vr,-
    1--,"',-n' int cTitrnh]I. i LllFtiI,. id Fial i i E1 .l' ri. 1 in tai.ii .d a f l f" 1;i t
    m )iIkt',, c uriir.]i r -;iir+f:.i,'e. (--'.'l l',4.p'..i'I ili~ l til. tlr,'. dli t.il!h fiu r't.e ret' cl I:b,
    q, th FiLii \ I i Kr1 I' 1 11 '- T. f iii. lnt c th
    s t iti f, SI
    ,lii i.tI lil l. t t i'rt r T e L tL lI "


    +fItt .: I 1r1i.i' 1f2 I\' i, \k r. i 'r iu 1i-,I l ii t- u i.I I, t--i I -irt !,, J i I A ;.. I ; iL,'
    bel':tC .% the S ift'_" C,'- -'r'k (-I,'<.s N \L..tti,,. .i:, (I,] 0 \ r' *i' r prI .pr -:' g the Ii.ii ,I -
    } P irn.' i i. r,' ,ij ,e, i n (1- -, I .! f 'I rivr \ l (z!i lli L i'', u4i f.

    1 .'rii ll;t i i. i T i 1 l ti ( 1' P[t' I.: .c'hn' raillw l '1 r : f'5, I n '. i : Ii th r f l1f'i I RN r ri]
    Is k'n;..,'l,. A ,ttiit'r p ipit-r. ',i, ir I i rs.I'J,,iia] d(-tta1 ls I, C:C' i i.",' irrt- ,. .f this
    ,f rirt it i,'ti "an1d Its f ,siii.V, 1, IIILrT _prt-'., t .. .

    Whereas ?.hc P'Furt iipr.*s- Z' rC./chip'IuS transition as dt(sciril f
    above is in-ot as ye.t subst ntiated, (ock's .Ip-Tr (Idos serve to p.o Tiit
    outi t l t-. M t crichijIpuS. (Lk w .s i ot .ipp,'r it il the C eat Plains seq nI-itICe
    bPelo,,' tlt R,1T11Tlin wa.1,iter i'morT t l ion. Schultz ( 193.S T. i l -. I ,
    Elias ( 19-42 ( C:idv 194 'i0 .1;. SchuIlty ;ind Stout i 194] 1, Sc'iulty .nal
    likh ] 1.9.1 1), aiId McKena.t i 19&65 all consid'Cr the Nlars-
    land Ior.IioI.tio. to l ~ t!t' h si fun,. ln i(ii o the Ilemingficid (Crotp Iil
    wv'esteri NtebraL.ska. ihus, if the concept of I hr Ilrihtiihrfordi.ati St ,.e
    Agct is l;isrd on the r-'ks I n d ii. ) sils 'nT ln stig t,.. H<1 ,i- B forI (1
    ( roi ip, its owcr l imi it sI C 111 nr ,(i.'Ti Ii r .' l i zed out .f the type
    Airca on the la,)1 ,f fossils th .t app-ear in the westt I e-ds of the NI. Irs-
    a1nd FOrn Iint1 Ict pu ;b' : on Lie other I(.-. nd ( ts itt .) oiP lar iii
    the st .~io I- ul- til it ]east as; 1iN in t1 sct-fm.l :.is the Ik 1mnin.w ater
    f1orn.tij iO whIj.ch iS v 0'oli ii.r tlian tl Nlarsnl .'t If t the crituerJa
    ()sbhorn : 1918~ : ad (C:.ok (1$)% 0 t) ftuso fo-r :-lcooircii M1"rfcipus.'
    are valid, ti:_n ,'id-iec.- from thei .stitatildraphiic occurred ce of thicst

    'C iok .nid o4t ii 19 3:3. list the f:.nia frinm b i 'e "'ppi i Harrisonri" 1 M\ar'ilti.d 1
    If Nei't:rska T 1'he r "Uppt.-r R'sch'iud" iTT Il b e in part ecit:valent to the Rui'ii i-
    water,


    2, '






    21O L T. LI TTN 'FLORIDA STAVE MNUSEL .I V, .. 14

    fossils indicates that Ithe practiC of c'',pl. ilig tphel first occu rre- r of
    ; /. t, ICt IS to 11111 t aLs o I t' I i.iili Un.iw.'dil. iS t 1t1 a s it
    i. b .;L'se t.l J tl '' ro.'ks iiIld fossils i- t[lI( l{('I2Iti!i -dll (;rull (. al .( t isl
    is the' sated ilte'tiol oa (t \Va l ; 12 i i tl.
    Aith 1 .'ii th1 ,'r'i ti r ci ) !iT., if it ; 1 -t;i t l i fr nf!! ifbl' / i, Ir' 1 i t 1t.'

    t if 'n tIrn U'Tur liwip)/) )."-curn i Tu tin} late' [ iljl-'n tordi n of the
    (Ii'..:t Pla ins it is N dCtn-'4ns it.I x 111 0. Iknow.n fr 1 -rni Li t .-t A rik.i cir'c.ir l.p sits in thdte reguio.. In poIt i, l actl it
    Otl.riplai r ID. .1 1 C' )Slcosely \V i 'I for () -ur uh1 I -S I II (t 1)i ftI 4 11 11 in(. i
    Vf.ti Tgip Icg, i. t.. t iI Iillnm gwatcr I. nd la ter.
    Urtil tli, tainnas cit1 tiw, '.IrsI1-3Ii I1: ltit mnTi .\a et<-Fr }t' i; titmis
    a iId] il o fu. n vll le n less r kniuO\ i L d FIill ll(d BoX BiliiC ( -
    posits bt-c ii0 t. i -ttT'i cr ksni \'il. thi L.:' cis d irit. siatui.i...nt I cdin rr.i ke
    1t o t tie pla 'c--nerit otf th ( ;. r iis ( .1 ut.ill int 1i titt, i. s te.nrn of
    their res p tix fu-!tr.i. it is pi'st- i:'r I1 arris n" and pri"-"'Sh ep
    (J-rIek. ( i tl.-i laid s of T' iturr .p. ( Iii 1 it. fv 1i- J 1 olf its f .au-1 I c' Iit l .~ tie nt .s. I H -!rix tha1 t :,if l' t <. i ti4l
    Cal.trvii ( 'lly i ii i .'Is ( ~ r)l lv \( .1>i! .itr ti an the Lower iL.Airi(son
    Sr (.1 is pro .b lv best c-or-rt-la cl. v,. I ti i[ I.t fLn.i!,-i Ir< n R iu iiii i ; dt Th sIrn l T ;- bT. I' said oi t ie [1 lorid -1 l'lIT- I.- S Flarill f. 1 i ', t-.'.-pt
    that it ai ppe'ars to 1" slight v i.iii 'r th i ti- (..i.ar -\ii (ullv.


    Bu 1 1 K.EVI I. II. FA ,uN.A


    il I1.111 na id a <-- l t r 1 sil I ;i }[ ;'i 1 vlzt1-'c'd (<.. on n'l i ts the Tho.n-i .s
    Fa iri. 'Tid Aill ,vOl 'lusr- cTrrelation with tile Oil r'\ :ii \1 l W:.iT
    FauLtlis of THInd. Tlis appears to be.c valid. The Burkt-ilie Pr.7p1-
    ti t 'ore -ra. fexanu., \wh "'h ich ;s I,,1if, 1 \,.i (.i. I tan tC Ga i 1 (s C ll x
    form closely r('s TiilIl.'. tilt." ".'h( .i. irri ". f1h ii tu. ( -aro. y tntY ffu-
    C" l '- d(r Pam1 S i W\ hit- a1 rd S F/q ?' ?f CTris Il r'a irefi \f\'Iltc i I'nrihfai ir-
    F t .s .pec-'n. r t. u froni W htil Burkt:- 'ill: e Fani ta aI-r S similar 1 lo F I < Sl .4 uh-
    ( tf heriu :. from t I h i T i as 1 ;.r li I ut. ti as' -',itm.C -id t .o a -e sptcit' s,
    F. tt'i.a7inr.s. Io' nit lurt..'s .'dul .." oC 'i -.1 is ii o li I ,sil i.:, as dht. :f,..ras/-.
    chiippus _'.unter li -.p-doi.. L? its.ri of Q. i-rn, 19,55 ).
    Its stratig rapiifc ()(f1lr:rIU 'Ti- ai. \'i -<- li. C '-nh'm in ,lo4 ,di'in __l\'ii
    Cull v Fau li .and th. ti and roel.ct, titv <' sa, I f ;, C llii. i 0 o 11 11 i-1 -t
    its fauuna ( Tal-, 2 ) ` i6 d Vtics that e B 1'rk evi FAUIa aI So should
    he sN.si.1.7I.d to the middle I -ernintlo-ir in the phyletic S.CL{unccCS of b)'th Pr('-.I /.tu're' lMand FioriHdatra--


    _ _ _____







    196 PATTON. TEX.AS ABRTOr)ACTYI.S 21]

    utilus between the (Garvini GCull and Burk-ville Faunis indicates that
    the time separating thet two fiaunas is of relatively short d rsltao-n.
    ()f the four biostratigraphlic units considered, in this paper, only
    the .Burkeville Fauna lacks came] material. This seemingly ano1nmalous
    situ tlioin iTiai b i part ~ 1 rcfr ctl 'iEl I tii ((.nCl i1parat ivelv poei orer state
    of k luwlic icd l o this t clua iD, .sMII1.i Irgii1 thi_ t S'pc it'iiS ily yv't
    found: or it may be-ir a"ttierfd ui to differ cti.-l! environmental sampling,
    wv'herebv onxi v' i'ertai ii halitat, a!ppIaircritlv iifh Voral tie to erN.1is, ,ere
    Filear Of tlil i .T fau ;, In any event, it is (difficult to conceive of entire caricl
    populationiis (ligratii fr l tloer Te(x.a(-s Coast cdiring the time the
    Burkevi le (dCr.'puits wcre acc-tumuirrating onl I to- return shortly there-
    after. IrPossibly local differential extinction was followed by a ,new
    spread, allhbiou i we h-ive 1 t i eCvideTi('c for tinis.
    Thle i t).1. ic inll the Buikcvillc F auidL ut otIier g1_ [aSLUI.l_1 ,-r .StJiln-
    T.i (1I dwellinIg IfonIts isilI as Amt ha/fpl--)n ai 1nd ti' oreod(]onts, arid the
    conriparativeilv di(m iihed hi orstf;. ftiia i.s si- ific.ant. [t is noteworthy
    thlit these form-s are present in relativeiv plentiful numbers in the
    faunas that bracket th c l rkt.vill Fauntr.t in tiiime. If the lack of cJamels
    is a result of li mitE.-d ihaitat sarrnpling, it is a most unifortlunate cir-
    C 1I in;tin with reg-ti'd to comprehending camel evolution in the
    Coastal Plain faiina-s, for witholut s11ch information it is virtually im-
    p1ossibl to spec.-cil.tec (11 thie iaiturlt of thlie riclationslisi) p itwe.'t the
    caieiIr.ls of the CGarvin GullWy "LFaunta anid thosc of thc Cold Spring. iThis

    '1ab]-c 26. A; iGE KAN E iS.r C-L.rt SPRING (; E.:I..A

    "'['i A' Rn A ri k:.i rr,- ;i.. T{ -nf'iI lf .l.rI ll l,.irst' vi i. t


    At I gdork

    TiU h.,ulcpus x x

    !A .'c pi,-a r,. .1fwjr tx x X
    I)i rrr.i.t hifTU P \
    Tich '.'Cra.s x
    Irr xx
    PTelicr'.1 crii X X

    P F'i, i t ,ras x x
    A.'e r r yit)us j N ,x


    ('ulirjpus x






    :12 B I_.LEV'I 11' ]1 (, i .'A T -: l _.'I SE Ul \I l. i.1

    be'ome.n s .[c)tL'.itiyV c-it t(a, fur 'nailp .. in 1 U- .itte lt t f-st.-ii -h
    the possible plihylognetic0 relio._-it ip o .- rala,"r/ ,a"In ., to A.qy-
    -aj 1 49' is

    COLD Sr,"N:cl F. rVA
    Th"ti Cold Spring FIa "II i.s eo,,t.iId bit Quinn i, 19... : to he a-.
    prox in teIl ot f Iate I Itmin lordian aek "T- ho rses of t.h C ol
    Spring scem ivT;rcilated to any f.ritis otter tiut those firor thl ( o Irr!I-tion. I H. the ( old Spri -ItIIna nIl. t. i,-f r .u 1c
    1'i; .I uai -' bIv th ie re lai h c v{ lll u I tior.);trW' It, sitl(0 (i t1](' fa(1'.1 it i)
    regArd to t I. BIfuI k vi ll itil which is c, rt.: in11)' older. aI .-I w Ihi
    og;it to the] L.apara (.i r uk i:.iit. whicl i.s ~e tiaili.I \ot l .i:r. IIl
    this _resp ct th C-.. ld Spriri fl,_ri a is c10arlv rnuch h ci sr ti o the
    rl1rk..'viic k thui i than to (1ic L ip ian (_n--i i-. t.,. ;-v,1 :.ie.: ns. t h, : :tp-
    p \ irn t L 1e]y .it rni: le 1 ioC I i (. Int. a, og."
    (0 i n ] 95.5: 74 ) fi.rthe r s its t;hat I "te iIli U't( iiN 'otInt OTT iTifts
    (of the (old Spring FaiLIIIa i appAtt--~u to have ro counterparts else
    where 'xc(pt for t ihe ('av I-rt inaLI erial i(. Ga.i:i In dt (1-1 o llii s1 150 .
    w\i, I.Ch 'orresponds Wit ith Cold Sprin I i-so(uar ;.s tlhe probostideans
    arTC COiernle." SvT ul lv kinollV to QUiil are Virtitalv iln',t'p;trahi frol it h.rn]is i ('tIri Tig
    fi. W 'I t 'in l41 L.i., Tichok i tl S ri lfis is I'port 'd li Sc .iltv.z and F-alk-
    c ibilch I 194 Ia t o n- -s 1 ii ib L,,e' S na ike (r-k iiatit i elts- Ir
    metir Of tlin r;W iis-ulna and Onfm'talfiriis t (] ( SYprinig A.p?/f7m 4lff.s
    cor respond to thc-isr, of A. a e.xanmi rao." lom il Barstow ftii;i oi (" :a -
    fc lia ( Davidson, 19. ; C( ld Spril S~ p c-iriiili aC.ssi 1 to !. onf
    rostrome ryx l icki corrn-tspo0d1 to tu, tp spc n. i] v'4s.. of that specie"-.-
    from the "uppermost" Santa Fh- bed; L; io-Plihcene of New M % --k:i_
    Frick, 1947. Simpson. 1950 ).
    None of the alloc'tho'i ,ous Cold Spni-g geflra except i" V rat h-
    :riurmn are reestrictet l4o tile -11ci1,ii1m frdia or h u o,. it ranLt,- cith'r
    inio the Bar sOViali or th.le C( lLrtC-i ti;tit. it' (ci' l ic] aid thl,- deer
    of thls Co-.ld. Spring arc adva -c.ed ovrc rclat'-td lit, I fl-min.lrdo rdi .an
    UtInIs in the Great Plains. G iJ; h l riium is not on fidenti- kIniow,'
    to appear in North Americ a-n -nia k \, -('lc- t hai;.u the Barstoian.
    VM1 -st od II remains v not bl(e, rcco\-t'rcd iron-) the lower Snake
    Creek deposits, but in the (rea! Plains aPppear for the- first time in
    thle Pawnee_, (_C-reek beds, which arc g.crierilly icgardcd as late Bar-
    stovin. Crutnphohruim from i tile (. alvex-t F"ormationi of %larv. iand
    is discussed 1b\ (IA .ix.i. aind (.5llns 9-0: ) i s b 'iii siKificait iin either
    of two ways": 'I" the pn'stnco:' Of ln,t stodl.il: js to b.I rgard.ed as placing




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