Front Cover
 Table of Contents
 Systematic index
 Cynarctoides sp.
 Indeterminate genus
 Age and correlation
 Literature cited
 Back Cover

Group Title: Bulletin of the Florida State Museum Biological sciences
Title: The Large mammals of the Buda local fauna (Arikareean
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00095833/00001
 Material Information
Title: The Large mammals of the Buda local fauna (Arikareean Alachua County, Florida)
Series Title: Bulletin - Florida State Museum ; volume 24, number 2
Physical Description: p. 123-173 : ill. ; 23 cm.
Language: English
Creator: Frailey, David
Donor: unknown ( endowment )
Publisher: Florida State Museum, University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1979
Copyright Date: 1979
Subject: Mammals, Fossil -- Florida -- Alachua County   ( lcsh )
Paleontology -- Miocene   ( lcsh )
Paleontology -- Florida -- Alachua County   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
Bibliography: Includes bibliographical references (p. 170-173).
General Note: Cover title.
Statement of Responsibility: David Frailey.
 Record Information
Bibliographic ID: UF00095833
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: oclc - 06423002
lccn - 80621193

Table of Contents
    Front Cover
        Page 121
        Page 122
    Table of Contents
        Page 123
        Page 124
        Page 125
        Page 126
        Page 127
    Systematic index
        Page 128
        Page 129
        Page 130
        Page 131
        Page 132
        Page 133
        Page 134
        Page 135
        Page 136
        Page 137
        Page 138
        Page 139
    Cynarctoides sp.
        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
    Indeterminate genus
        Page 154
        Page 155
        Page 156
        Page 157
        Page 158
        Page 159
        Page 160
        Page 161
        Page 162
        Page 163
        Page 164
        Page 165
    Age and correlation
        Page 166
        Page 167
        Page 168
        Page 169
    Literature cited
        Page 170
        Page 171
        Page 172
        Page 173
        Page 174
    Back Cover
        Page 175
Full Text


of the
Biological Sciences

Volume 24 1979 Number 2





SCIENCES, are published at irregular intervals. Volumes contain about 300 pages and
are not necessarily completed in any one calendar year.


RHODA J. RYBAK, Managing Editor

Consultants for this issue:



Communications concerning purchase or exchange of the publications and all
manuscripts should be addressed to: Managing Editor, Bulletin; Florida State
Museum; University of Florida; Gainesville, Florida 32611.

Copyright 1979 by the Florida State Museum of the University of Florida.

Publication date: December 17, 1979

This public document was promulgated at an annual cost of
$1,851.30, or $1.851 per copy. It makes available to libraries,
scholars, and all interested persons the results of researches in
the natural sciences, emphasizing the circum-Caribbean region.

Price: $2.00



SYNOPSIS: The large mammals of the Buda Local Fauna are discussed. Two new
species are described, Daphoenodon notionastes and Bassariscops achoros, and a new
genus of Camelidae is recognized but not named. Nanotragulus lulli and N. intermedius
are synonymized with N. loomisi. Additional taxa included are Cynarctoides sp.,
Mustelidae gen. et sp. indet., Nimravinae, gen. et sp. indet., Cynorca sp.,
Phenacocoelinae gen. et sp. indet., Camelidae gen et sp. indet., Anchitheriinae gen. et
sp. indet., and Moropus sp. Biostratigraphic correlation indicates that the Buda Local
Fauna is Arikareean in age.


INTRODUCTION ........................................................ 124
ACKNOWLEDGEMENTS ......................................... ....... 127
SYSTEMATIC INDEX ............... .. ......................... ......... 128
AMPHICYONIDAE .................... .... ..................... ........ 128
Daphoenodon notionastes new species ................................ 128
CANIDAE ................... ................................ ........ 134
Bassariscops achoros new species ....... .......... ..................... 134
Cynarctoides sp ................................... ............ 140
M USTELIDAE ...................................... ............... 142
FELIDAE ................... . ................... ............. 143
N IM RAV INAE ................................................ .. 143
E QUIDAE ...................................... ............ ...... . 144
ANCHITHERIINAE ................................................ 144
C HALICOTHERIIDAE .................................................. 144
Moropus sp....................................................... 144
TAYASSUIDAE ................... ...... ...... ................... . 149
Cynorca sp. ...................................................... 149
M ERYCOIDODONTIDAE .................... ............... ... ...... 150
PHENACOCOELINAE ..................... ................... .......... 150
CAMELIDAE ................................. ......... 151
NEW GENUS AND SPECIES ........ ......... ................... ....... 151
INDETERMINATE GENUS ............................................ 154
HYPERTRAGULIDAE ........................ ......................... 156
Nanotragulus loomisi .............................................. 156
AGE AND CORRELATION .................. ................... .......... 166
ZOOGEOGRAPHY ..................................... ......... ........ 168
S U M M A RY .................................................... ....... 169
LITERATURE CITED ................... ...... ...................... 170

'Department of Systematics and Ecology and Museum of Natural History, University of Kansas, Lawrence, Kansas

FRAILEY, DAVID. 1979. The Large mammals of the Buda Local Fauna (Arikareean,
Alachua County, Florida). Bull. Florida State Mus., Biol. Sci. 24 (2):123-173.



The early and mid-Tertiary history of land mammals is very sparsely
recorded in eastern North America. The broad sheets of continental
sediments, which so beautifully preserved the successive faunas of
western North America, have no counterparts in the East. Of all the
eastern states, Florida has yielded the best collections of Oligocene
and Miocene land mammals. This fortunate occurrence is due in great
part to the widespread carbonate deposits that underlie much of the
Florida peninsula. These marine carbonate deposits of the early Ter-
tiary became riddled with solution cavities later in the Tertiary and
were subsequently filled with plastic sediments. Such sinkhole
deposits are small, randomly scattered, yield little stratigraphy, and
are often destroyed before their presence is discovered. They are form-
ing today in Florida in a wide variety of habitats ranging from swamp
to upland areas (Harper 1914) and presumably did the same in the
past. Sinkholes are natural traps, often attract animals because of the
water they sometimes contain, and involve little transportation of an
animal's remains after its death. Preservation is often exceptionally
good. Unfortunately, fewer than a dozen mid-Tertiary sinks are known
that contain vertebrate remains. Of these, most produced only a hand-
ful of fossils, the rest of the fossils having been lost during the mining
operations that first uncovered the sites. The three most prolific mid-
Tertiary sites have been 1-75 of Whitneyan late Oligocene age (Patton
1969a); Buda of late Arikareean very early Miocene age (this paper),
and Thomas Farm of Hemingfordian early Miocene age (Simpson
1932; White 1942; Olsen 1962; Patton 1967). They provide most of
what is known about the Oligocene and Miocene land animals of
Florida and, for that matter, of eastern North America.
Several fossil vertebrate local faunas in Florida have been described
as Arikareean (very early Miocene) in age: Thomas Farm; Griscom
Plantation (Simpson 1932); Seaboard Air Line Railroad Company,
Tallahassee (Olsen 1964); Franklin Phosphate Pit No. 2, Newberry
(Simpson 1932); Brooksville (Patton 1967); and SB-1A, near Live Oak
(Frailey 1978). Two of these sites, Griscom and Seaboard, are small
and dated primarily on their faunal relationships to the better known
Thomas Farm Local Fauna. Studies of the fauna (Patton 1967, 1969b;
Tedford and Frailey 1976) now date Thomas Farm as early Hemingfor-
dian in age. On the basis of close resemblance to Thomas Farm,
Griscom and Seaboard may also be transferred to the Hemingfordian
faunal age. Franklin Phosphate Pit No. 2, Brooksville, and SB-1A then
remain as the only fossil faunas in Florida still thought to be
Arikareean in age. Another site containing the Buda Local Fauna is

Vol. 24, No. 2


the focus of the present study and is the largest Arikareean sample
known from eastern North America.
In February, 1965, the Buda Local Fauna was discovered in NW /4,
NW 4, S 32, T 8s, R 17E, near the small town of Buda, Alachua Coun-
ty, Florida (see Fig. 1), and extensively collected by S. D. Webb, N.
Tessman, J. S. Waldrop, and E. Kayworth during field reconnaissance
for the Florida State Museum. Subsequent collections were carried out
by T. H. Patton, J. G. Klein, and Fred Dixon of the Florida State
Museum. The fauna was discovered in the Buda Mine where limestone
strip mining had cut into a fossiliferous clay-filled sinkhole. The
sinkhole was situated within 10 yd of the south wall of the limestone
mine and was not completely destroyed. It consisted of three shallow
vertical chambers of from 2 to 8 ft in diameter which presumably had
shared a common opening. The fossiliferous clays were oxidized only
where they were in contact with the limestone wall.
Several fossils were found in place during the initial collecting.
These include parts of Daphoenodon notionastes n. sp., the uniden-
tifiable mustelid; Nanotragulus loomisi, the oreodont; an unnamed
new genus of camel; a larger unidentifiable camel; the horse; and the
chalicothere. The majority of the fossils were recovered by searching
the spoil banks and by collecting pockets of clay which were later
broken down and screened. The distribution of fossils in the spoil
banks, as recalled by those who collected the locality, was not random
(see Fig. 1). The small mammals were found in a clayey sand, in con-
trast to the clay sediments associated with the other mammals.
This study is limited to the 12 larger mammals in the Buda Local
Fauna. These and the taxa of Brooksville, Franklin Phosphate Pit No.
2, and SB-1A are listed in Table 1. The hedgehog, Amphechinus sp.,
has been described by Rich and Patton (1975). The other small mam-
mals, reptiles, amphibians, and fish will be reported on at a later date.
The majority of identifiable fossils in this fauna consists of isolated
teeth. This imposes certain limitations on the interpretations that can
be made. I have hesitated to recognize isolated teeth as holotypes, but
where composite samples are adequate I feel that the differences will
remain readily apparent and that the uniqueness of the sample
justifies this course.
The specimens comprising the Buda Local Fauna are curated in the
Collection of Fossil Vertebrates, Florida State Museum, University of
Florida (abbreviated UF). Other abbreviations and symbols are as
follows: AMNH, American Museum of Natural History; CM, Carnegie
Museum; F:AM, Frick American Mammals, American Museum of
Natural History; FGS, Florida Geological Survey Collection (now part
of the Florida State Museum Collection); HC, Harold Cook Collection



FIGURE 1.-The Buda Locality. General location and distribution of some of the fossils
over the spoil piles.

(now part of the American Museum of Natural History Collection);
MCZ, Museum of Comparative Zoology, Harvard University; PU,
Princeton University; SDSM, South Dakota School of Mines; UC,
University of Chicago; UCMP, University of California, Museum of
Paleontology; YPM, Yale Peabody Museum; ( ), approximate
measurement; N, number of specimens in sample; OR, observed range;
X, mean; SD, standard deviation; CV, coefficient of variation. All

Vol. 24, No. 2


measurements are in millimeters. The drawings were made with a
camera lucida microscope.



Amphechinus sp.
Daphoenodon notionastes n. sp.
Bassariscops achoros n. sp.
Cynarctoides sp.
Mustelidae gen. et sp. indet.
Nimravinae gen. et sp. indet.
Anchitheriinae gen. et sp. indet.
Moropus sp.
Cynorca sp.
Phenacocoelinae gen. et sp. indet.
Camelidae n. gen. et sp.
Camelidae gen. et sp. indet.
Nanotragulus loomisi

(modified from Patton 1967)

Daphoenodon notionastes n. sp.
Rhinocerotidae gen. et sp. indet.
Tapiridae gen. et sp. indet.
Merycoidodontidae gen. et sp. indet.

Franklin Phosphate Pit No. 2
(modified from Simpson 1930)

Daphoenodon notionastes
Anchitheriinae gen. et sp. indet.
Rhinocerotidae gen. et sp. indet.
Entelodontidae, ?Daedon
Camelidae, ?Oxydactylus
Blastomericinae or the unnamed new
camelid from Buda'

(from Frailey 1978)

Protosciurus sp.
Mammacyon cf. obtusidens
Canidae gen. et sp. indet.
Phlaocyon sp.
Paroligobunis frazieri
Carnivora gen. et sp. indet.
Anchitheriinae gen. et sp. indet.
Nothokemas waldropi

'This identification is based on three upper molars, now lost, but two of which were figured by Simpson (1930); he
referred these teeth to cf Blastomervx. The features of these teeth, including those which bothered Simpson, can be
seen in the upper molars of the new genus from Buda.

I wish to thank those persons who aided and encouraged me during the writing of this
paper. Principle among these are Thomas H. Patton, S. David Webb, and H. K. Brooks
of the University of Florida. In addition, I wish to acknowledge Malcolm McKenna,
Richard Tedford, Beryl Taylor, and Ted Galusha of the American Museum of Natural
History, Larry D. Martin of the University of Kansas, Robert M. Hunt, Jr., of the
University of Nebraska, and Margery C. Coombs of the University of Massachusetts
who donated time and knowledge to this project. Colleagues of mine, Jean Klein, John
Waldrop, Michael Hansinger, John Meeder, and Ronald Wolff, provided invaluable
companionship and discussion during the course of this project. Edythe Humphries,
Kay Purinton, Nancy Halliday, Deb Bennett, Lorraine Meeker, and Chester Tarka lent
their talents and advice in the preparation of the photographs and illustrations. A
special note of appreciation is extended to Philip Bjork of the South Dakota School of
Mines, Bryan Patterson of Harvard University, and Mary Ann Turner of Yale Univer-
sity for lending specimens under their care.


Daphoenodon notionastes new species
ETYMOLOGY.-notios (Gr.): Southern; nastes (Gr.): Inhabitant.
HOLOTYPE.-UF 16965, M,.
REFERRED MATERIAL.-UF 16936, 17 incisors; UF 16938, 3 upper premolars; UF
16906, P4; UF 18499, P': UF 16944, M2: UF 16921, 3 C,'s; UF 16940, 5 P,'s; UF 16968,
P,; UF 16901, 3 M, talonids; UF 16905, M2; UF 18356, M3; UF 17001, maxillary frag-
ment with alveoli for M'-2; UF 16910, edentulous mandible; UF 16970, 3 fragments of
mandibles, 1 with P,; and UF 16904, 16909, 16912, 16917, 16926, 16948, 16993, 16995,
16996, 17009-17011, 18338-18355, various postcranial elements.
COMPARATIVE MATERIAL EXAMINED.-Daphoenodon superbus: CM 1589A, 1589B,
1589D, 2774, 2199; AMNH 81003, 81025; FGS 1213, 1214; PU 11554. Pericyon socialis:
YPM 12715, holotype.
DIAGNOSIs.-Daphoenodon notionastes is the smallest species of
Daphoenodon. It is about 20% smaller than D. superbus and about
45% smaller than D. robustum2. The Mi is low crowned and widest at
the protoconid. The rim of the talonid is incomplete, forming a depres-
sion between the entoconid and the metaconid. The talonids of M, and
M, are noticeably smaller (in length and width) than their respective
trigonids. The mandible is slender and very Canis-like in appearance.
The depth of the mandible beneath M, is less than 11/2 times the length
of Mi.

UPPER DENTITION.-The P' (Fig. 2, E-F) of Daphoenodon no-
tionastes is smaller than that of D. superbus, but not separable other-
wise. A broken P4 (UF 18499) is smaller than the only complete P4 (UF
16906) and indicates that some individuals were more than 20%
smaller than D. superbus.
No M' of Daphoenodon was found at Buda.
The M2 of D. notionastes is essentially flat, the paracone being only
slightly raised in contrast to the elevated paracone of M2 in D. super-
bus, which in the latter produces a gentle concave curve when the
crown is viewed anteroposteriorly. Few M2's are housed in collections,
and the differences listed here may not be diagnostic.

'In this usage I am following Hunt (1972).
2Daphoenodon robustum was originally described as Borocyon robustum by Peterson 1910. This species was trans-
ferred to Daphoenodon by Hunt (1971), and Borocyon was dropped. Hunt also synonymized D. niobrarensis Loomis
1936 and D. periculosis Cook 1909 with this species. Thus, Daphoenodon superbus and D. robustum are the only
other valid species of this genus in the literature.

Vol. 24, No. 2




0 5cm

FIGURE 2.-Daphoenodon notionastes n. sp. A and C) UF 16905, M,, lingual and oc-
clusal views; B and D) UF 16965, holotype, M,, lingual and occlusal views; E and F) UF
16906, P4, lingual and occlusal views; G) UF 16910, left ramus. Natural size.


TABLE 2. COMPARATIVE MEASUREMENTS OF Daphoenodon notionastes AND D. super-

D. superbus

P' (length x width)

M2 (length x width)
P, (length x width)

M,(length x width)

M, talonid (length x width)

M. (length x width)

M, talonid (length x width)
Depth of mandible at M,

Greatest length
Anteroposterior diameter of head
Transverse diameter of head at
the tuberosities
Greatest transverse diameter of

Greatest anteroposterior diameter
of distal end
Anteroposterior diameter at
coronoid process
Transverse diameter at coronoid
Width of ilium at great sacro-
sciatic notch
Anteroposterior diameter of

Vertical diameter of acetabulum


22.0 x 13.6 (CM 1589A)
23.4 x 13.9 (CM 2774)
19.9 x 13.4 (CM 1589A)
18.4 x 8.6 (CM 1589A)

21.7 x 10.0 (PU 11554)
23.5 x 11.0 (CM 2774)
25.5 x 11.5 (CM 1589A)
8.8 x 11.0 (CM 1589A)
7.0 x 9.4 (PU 11554)
12.2 x 8.8 (PU 11554)
16.6 x 9.6 (AMNH
6.6 x 8.1 (PU 11554)
29.8 (CM 1589A)
36.5 (CM 2774)
37.3 (PU 11554)
39.3 (AMNH 81003)










D. notionastes

20.0 x 11.3 (UF 16906)

14.2 x 8.4 (UF 16944)
14.9 x 6.0 (UF 16970)
15.6 x 6.8 (UF 16968)
21.6 x 10.6 (UF 16965)

8.1 x 9.6 (UF 16965)

12.3 x 8.7 (UF 16905)

6.5 x 7.6 (UF 16905)
28.4 (UF 16910)
30.2 (UF 16997)

162 (UF 16948)
40 (UF 16948)

32 (UF 16948)

28 (UF 16948)
34 (UF 18347)

33 (UF 18347)

25 (UF 18349)

22 (UF 18349)

24 (UF 18348)

20 (UF 18348)
21 (UF 18348)
27 (UF 18348)
20 (UF 18348)
22 (UF 18348)
25 (UF 18348)

(176) (UF 18352)

Vol. 24, No. 2




D. superbus

Anteroposterior diameter at lower
end of cnemial crest

Transverse diameter at lower end
of cnemial crest

Greatest anteroposterior diameter
of distal end
Greatest transverse diameter of
distal end

Greatest length

Greatest transverse diameter

Length of tuber from susten-
taculum to free end

Greatest height

Transverse diameter of trochlea

Transverse diameter of head

Metacarpal II
Metacarpal IV
Metacarpal V
Metatarsal I

Metatarsal III
Metatarsal IV

D. notionastes

19 (UF 18352)
21 (UF 18352)
22 (UF 18352)

15 (UF 18352)
17 (UF 18352)
18 (UF 18352)

20 (UF 18352)

(29) (UF 18352)
28 (UF 18352)

48 (UF 18354)
58 (UF 16912)
25 (UF 18354)
31 (UF 16912)
32 (UF 16912)

28 (UF 18354)
34 (UF 16912)
36 (UF 16912)

32 (UF 18353)
36 (UF 16995)
39 (UF 16995)
15 (UF 18353)
18 (UF 16995)
19 (UF 16995)
16 (UF 18353)
18 (UF 16995)
19 (UF 16995)

38 (UF 18339)

49 (UF 18340)

44 (UF 18341)

28 (UF 18342)
30 (UF 18342)
33 (UF 18342)
35 (UF 18342)

70 61 (UF 16992)

73 63 (UF 18337)



LOWER DENTITION.-Except for its smaller size, P4 of Daphoenodon
notionastes is inseparable from that of D. superbus.
A low rim, which does not extend anteriorly to the entoconid, closes
the posterior boundary of the talonid of M, (Fig. 2 B, D) of D. no-
tionastes. In D. superbus this rim continues beyond the entoconid to
abut against the metaconid. In D. superbus wear produces a notch
anterior to the entoconid similar to the unworn condition of D. no-
tionastes. The talonid of M, in D. notionastes is shorter and narrower
than the trigonid. The metaconid of the M, of D. notionastes has a
slight posterior slant, unlike the vertical metaconid of D. superbus.
The holotype of Daphoenodon notionastes (UF 16965) is the largest
M, (and the only complete one) in the sample. Measuring 21.6 x 10.6,
length x width, it is at the low end of the measured range of D. super-
bus (21.7 x 10.0-25.5 x 11.5), a remarkable feature when one considers
the difference in sturdiness between the mandibles of D. superbus and
D. notionastes as discussed in the following section.
The M2 (Fig. 2 A, C) of D. notionastes has a talonid that, even more
than that of M,, is decidedly shorter and narrower than the trigonid.
This stands in contrast to the wide, elongate talonid of D. superbus
that equals the trigonid in occlusal area and gives the Ms of that
species a rectangular occlusal outline.
MANDIBLE.-The mandible (Fig. 2G) of Daphoenodon notionastes
is much less robust than the mandible of D. superbus. The massateric
fossa is shallower than in D. superbus, and the line that indicates the
separation of the attachment areas for the temporal and masseter
muscles is faint, indicating weaker jaw musculature for D. no-
tionastes. The depths of the mandibles referred to D. notionastes,
measured at M,, are 28.4 and 30.2. This is comparable to the size of a
juvenile D. superbus mandible (CM 1589A), which measures 29.8, but
is definitely smaller than the 36.5-39.3 range measured for adult man-
POSTCRANIAL SKELETON.-Other than the smaller size, most of the
postcranial elements referred to Daphoenodon notionastes are vir-
tually indistinguishable from those of D. superbus (CM 1589B). Some
elements can be grouped into two size classes and may represent sex-
ual dimorphism, but others present a more continuous range of sizes.
All are generally smaller than the corresponding elements of D. super-
bus. Two humeri (UF 16996, 18347), while essentially amphicyonid in
their features, are much larger than other humeri of D. notionastes
(UF 16948) from Buda. They also have an oval entepicondylar foramen
instead of the elongate, slitlike foramen more typical of amphicyonids.
No entepicondylar foramina are preserved on the other, smaller humeri
referred to D. notionastes. It therefore cannot be determined whether

Vol. 24, No. 2


an oval foramen is a variable feature of D. notionastes or that these
two humeri represent a larger amphicyonine that is presently not
recognized in the fauna.
Other than size, no other significant differences exist between the
postcranial skeleton of Daphoenodon notionastes and D. superbus.
Measurements of postcranial elements of D. notionastes are compared
in Table 2 to measurements listed by Peterson (1910) for D. superbus.

Daphoenodon notionastes has been collected in two other local
faunas of Florida (Brooksville and Franklin Phosphate Pit No. 2), but
its affinities have previously been unclear because of insufficient
material. With the more complete sample from Buda, the recognition
of D. notionastes in the Brooksville and Franklin Phosphate Pit No. 2
local faunas is possible.
An M, and a mandibular fragment (FGS V-1213 and 1214) were
found in Franklin Phosphate Pit No. 2 in Alachua County. These were
originally referred to Mesocyon iamonensis by Simpson (1929), who
later (Simpson 1932) transferred them to ?Temnocyon. Olsen (1958),
working without the diagnostic M2, felt certain that the specimens
were indeed Temnocyon. Hunt (1971) recognized the features of
Daphoenodon in the tooth and referred the specimens to D. superbus.
FGS V-1213 is slightly smaller than UF 16965 and measures 21.4 x
9.8. UF 16965 is the largest Mi in the Buda sample, and the size dif-
ference between this tooth and that from Franklin Phosphate Pit No. 2
does not seem to be critical. These two M,'s also have the posterior tilt
of the metaconid, absence of the talonid rim between the entoconid and
the metaconid, and the same narrowing of the talonid. FGS V-1213
does have an indistinct entoconid, unlike UF 16965, but this may be
variable within the species. The similarities override the small dif-
ferences and lead me to conclude that FGS V-1213 should be referred
to the new species from Buda.
Among the fossils from Brooksville, Hernando County, Florida, is
an isolated M, (UF 449) that looks much like that of D. notionastes,
although it is smaller (19.3 x 9.4). This M, has a similarly incomplete
talonid rim and posteriorly tilting metaconid that unites the
Daphoenodon from Buda with that from Franklin Phosphate Pit No. 2
and has the indistinct entoconid of the latter. The Brooksville M, is
stockier and has a relatively large, i.e. unreduced, talonid. The talonid
is the widest part of the tooth and very similar to D. superbus (or a
stocky Daphoenus) in this respect. An isolated M' from Brooksville
looks like a miniature of the M' of D. superbus and measures 22.0 x
16.1. Although the M, from Brooksville has a relatively larger talonid



than the M, from Buda, the size of the tooth and other diagnostic land-
marks are most in keeping with the new species of Daphoenodon
described in this paper.
The greater size and general robustness of the teeth and postcranial
skeleton place this species nearer to Daphoenodon than to Daphoenus,
its probable Oligocene precursor. Furthermore, the morphology of
Daphoenodon notionastes closely resembles that of D. superbus from
western Nebraska.
Daphoenodon notionastes of Florida represents a southeastern
group of Daphoenodon that apparently was distinct from its earliest
occurrence (Brooksville). Daphoenodon notionastes was smaller than
other species of Daphoenodon, especially D. robustum. Unlike most
amphicyonids, including D. superbus, Daphoenodon notionastes deem-
phasized the crushing function of its dentition by reducing the
talonids of its lower molars. The mandible was shallower and contrasts
with the massive bear-like mandibles of other amphicyonids.

Bassariscops achoros new species
ETYMOLOGY.-achoros (Gr.): Wanderer.
HOLOTYPE.-UF 18389, P'
REFERRED MATERIAL.-UF 16933, 15 incisors; UF 22778, 4 P''s; UF 16969, 2 M"s;
UF 18501, M'; UF 16963, 5 M"s; UF 18403, M' ; UF 16961, 10 lower premolars; UF
18412, DP,; UF 18411, 2 DP, trigonids; UF 18410, Dp, talonid; UF 16989, M,; UF
18390, 3 M, talonids; UF 16964, 2 M,'s; UF 16962, M,; UF 16991, 2 lower mandibular
rami, 1 with M, talonid and M,; UF 18391, fragment of premaxilla; UF 18392, fragment
of maxilla; UF 18397, 3 petrosals; and UF 16976, 16978, 16990, 18393-18396,
18398-18402, 18416, 18502, 19312, various postcranial elements.
COMPARATIVE MATERIAL EXAMINED.-Bassariscops willistoni, CM 11332, holotype,
CM 11333, 11334 (casts in AMNH); Phlaocyon leucosteus, AMNH 8768, holotype;
Nothocyon annectans, CM 1602, holotype (cast in AMNH); Nothocyon geismarianus,
UCMP 90, Nothocyon latidens, AMNH 6896, holotype, Nothocyon lemur, AMNH
6888, holotype.
DIAGNOSIS.-P4 is about 20% smaller than that of Bassariscops
willistoni, the only other species of Bassariscops. P4 has a smaller
hypocone than B. willistoni and lacks the strong labial cingulum of
that species. Upper molars are narrower and more angular in outline
than those of B. willistoni. M1 of B. achoros has the metaconid placed
farther posterolabially than is seen in B. willistoni. The hypoconid and
entoconid of M1 are equal in height, unlike that of B. willistoni which
has a taller hypoconid.

Vol. 24, No. 2


B. achoros B. willistoni
P. 7.8 x 4.3 (UF 18389) 9.6 x 5.4 (CM 11332)
8.0 x 4.7 (UF 22778)
M' 6.5 x 7.9 (UF 18403) 5.4 x (8.5) (CM 11332)
M' 4.1 x 6.2 (UF 18403) 3.3 x 6.5 (CM 11332)
M, 8.6 x 3.6 (UF 16989) 7.6 x 3.3 (CM 11334)

UPPER DENTITION.-P4 (Figs. 3A, 4A-B) of Bassariscops achoros is
very much like that of B. willistoni except for its smaller size. The P' of
B. willistoni has a stronger external cingulum as compared to the
barely discernible external cingulum of B. achoros. The hypocone is
slightly smaller than that of B. willistoni. The overall impression is of
a more sectorial tooth in B. achoros.
M'-2 (Figs. 3B, 4C) are heavily worn, and the labial margins are
broken on the holotype of B. willistoni. In general outline and shape
these teeth have some of the appearance of M'-2 on Phlaocyon
leucosteus, also heavily worn. This similarity may be effected in part
by wear and breakage, but it appears to be genuine. The angular, squat
(anteroposteriorly expanded) upper molars of B. achoros contrast with
the smooth outline and narrow upper molars of B. willistoni. Rem-
nants of cusps on M'- of B. willistoni indicate the same placement and
prominence as in B. achoros.
As M'-2 are so heavily worn on the holotype of B. willistoni, M'-2 of
B. achoros will be described in detail. M' of B. achoros is trapezoidal
and variably cusped, but usually has five cusps. The paracone and
metacone are of nearly equal size and together form the widest part of
the tooth. The paracone has a labial deflection similar to that seen in
Parictis and some other mustelids. The protocone is twice as large as
the metaconule, but only slightly more lingually placed. The
paraconule is variable in size and may be absent altogether. An addi-
tional cusp, as large as the metaconule, occurs between the metaconule
and the metacone on one specimen (UF 18501). The internal cingulum
begins anteriorly at the paracone as a thin line and lingually becomes a
heavy cingulum that rises to a single definite cusp, the hypocone, just
posterolingually to the metaconule before decreasing again to a thin
cingulum that terminates posteriorly at the metacone. The hypocone
is equal in height to the protocone, and both are of greater height than
the metaconule. The position of the anterior root beneath the paracone
in M' of B. willistoni is comparable to that seen in B. achoros. The M'





S0 5mm



C D c
FIGURE 3..-Bassariscops achoros n. sp. A) UF 18389, P', holotype; B) UF 18403, M'-2;
C) UF 16961, P4; D) UF 16989, M,; E) UF 16691, M,. A and B are occlusal views; C, D,
and E are shown in occlusal and labial views.
of B. willistoni looks like a lingually stretched version of the stockier
M' of B. achoros.
In the M2 of B. achoros the paracone is much larger than the minute
metacone. There is a strong labial cingulum on the paracone (the
parastyle) and no labial cingulum on the metacone. The paraconule and

Vol. 24, No. 2




0 2cm

0 2cm

FIGURE 4.-Bassariscops achoros n. sp. A) UF 18389, P', holotype, lingual view; B) UF
18389, labial view; C) UF 18403, M'-2, labial view; D) UF 16991, Right mandibular
ramus with M,.

metaconule may be absent and are variable in size when present.
Relative to the protocone, the metaconule of M2 is much smaller than
in M'. The internal cingulum is heavy, as in M'; the only definite cusp
on this cingulum is the posterolingual hypocone. The M2 varies in size
but is relatively large in this species.
LOWER DENTITION.-As no P, of B. willistoni has yet been found,
this is the first description of a Bassariscops premolar. In P, (Fig. 3C)
the protoconid is placed forward in the tooth, almost directly over the
anterior root. The posterior accessory cusp is labial to the midline of
the tooth; the posterior margin of the tooth is upturned. P, of B.
achoros is similar to the P,'s of many other small carnivores.
The M,'s referred to B. achoros differ considerably from that re-
ferred to B. willistoni by Peterson (1928). In B. achoros, the metaconid
of M, (Fig. 3D) lies posterolabially to the protoconid. The entoconid
and hypoconid are equal in height and surround a basined talonid. The
hypoconulid is indistinct. It agrees with B. willistoni in having a thin
protoconid with small anterior and posterior blades projecting from
the central cone. The M,'s of both B. achoros and B. willistoni have
deep carnassial notches. Both Bassaricops achoros and B. willistoni



have a small interoconid on the M1. Another tubercle, little more than
a line on B. achoros and B. willistoni, extends up the posterolabial
margin of the protoconid to the same height as the metaconid.
Whereas M1 of B. willistoni has a hypoconid that is slightly higher
than the entoconid, surrounding a large talonid basin, these cusps are
of nearly equal size in B. achoros. The talonid of M, in B. achoros has a
straight posterior margin as in B. willistoni.
The M2's of B. achoros and B. willistoni are essentially the same.
Each has a small paraconid, the protoconid and metaconid are of equal
size, and the metaconid sits slightly posterior to the protoconid. The
hypoconid is prominent. There is a large anterolabial cingulum on the
M2 of each species. This cingulum in B. achoros may be interrupted at
the protoconid (Fig. 3E) or may be complete and form a small ac-
cessory cusp posterolabially to the protoconid.
M3 of B. willistoni is unknown. In B. achoros this tooth is small,
single-rooted, and has indistinct cusps.
MANDIBLE.-Mandibular specimens referred to B. achoros and B.
willistoni consist of broken rami only. The mandible of B. achoros (Fig.
4D) is deepest beneath M2 and becomes markedly narrower anteriorly,
much like that of Cynarctoides. It differs from the more uniformly
deep mandibles of Nothocyon and B. willistoni.
POSTCRANIAL SKELETON.-The postcranial skeleton of B. willistoni
is unknown. Numerous fragmented postcranial elements of a small
carnivore in the Buda Local Fauna are attributed to B. achoros.
The humeri assigned to B. achoros (UF 18393, 18395) have
transversely elongated condyles, small subcircular entepicondylar
foramina, and small and gently curving supracondyloid crests which
resemble the humeri of the Procyonidae more than those of any other
canoid family.
The astragalus is more canid-like in its very anterior-facing head
and its deep trochlea. The astragalus is flattened like that of the
astragali of mustelids.
The calcaneum bears more definitely canid features. It is narrow,
the anterior part is long, and the lesser process is placed well pos-
teriorly. The posterior part of the calcaneum equals the anterior part in
length. The posterior part has its greatest height at the sustentacular
facet and gradually decreases in height posteriorly. The sustentacular
facet is procyonid-like, being less sharply curved than in canids.

The only previously known species of Bassariscops, B. willistoni,
was found in the lower part of the Brown's Park Formation
(?Arikareean) of Colorado (Peterson 1928). If B. achoros is an earlier

Vol. 24, No. 2


occurring species than B. willistoni, changes that could be interpreted
as evolutionary trends are the labiolingual elongation of the upper
molars and the slight increase in the size of P4 from B. achoros.
Bassariscops willistoni was originally included as a new species of
Phlaocyon by Peterson (1924). Phlaocyon at that time was regarded as
a procyonid, but now, mainly as a result of Hough's (1944, 1948)
studies of the auditory region of the Canoidea, Phlaocyon is generally
accepted as a canid. With the familial designation of Bassariscops in
doubt, I compared three periotic bones (UF 18397) that were referred
to Bassariscops achoros with the auditory regions of Canis familiaris
(UF 5680), Procyon lotor (UF uncatalogued), Bassariscus astutus (UF
7865), Mustela frenata (UF uncatalogued), and Taxidea taxus (UF
The mustelids were quickly eliminated. Segall (1943) describes
several characteristics which separate mustelids from other canoids,
only two of which (the stylomastoid foramen and the tympanic cavity)
could be used on the isolated periotic bones from Buda. These,
however, were sufficient. The stylomastoid foramen (here referring to
the stylomastoid foramen definitivum of some authors) is almost com-
pletely surrounded by the bulla in mustelids, unlike the equal par-
ticipation of the bulla and the mastoid portion of the temporal bone in
forming the stylomastoid foramen of Bassariscops. More conclusively,
the tympanic cavity in mustelids extends well posteriorly to the pro-
montory and only slightly anteriorly; the reverse expression of the
tympanic cavity occurs in Bassariscops.
Hough (1948) observed that the reflected margin of the bulla covers
only the base of the promontorium in the Procyonidae, but overlaps
the promontorium extensively in the Canidae. In Bassariscops the
characteristic rugosity indicating the area of contact with the tym-
panic bulla extends to the posterior lip of the fenestra cochlea, as in
Canis, but unlike either Procyon or Bassariscus.
The shape of the definitive stylomastoid foramen is useful to
separate procyonids from canids. The medial terminus of the definitive
stylomastoid foramen (inside the bulla) in Bassariscops is slitlike, as in
Canis, and differs from the round definitive stylomastoid foramina of
both Procyon and Bassariscus.
Another characteristic mentioned but not listed as diagnostic by
Hough (1948) is the grooving of the mastoid process for the facial
nerve. This grooving is shallow in Procyon and Bassariscus but deep in
Canis and Bassariscops. Other similarities between Canis and
Bassariscops include the long anterior process of the petrosal which
slopes forward to meet the basisphenoid, unlike the shorter anterior
margin in procyonids. The proximity of the canal for the facial nerve to


the suture between the temporal bone and the occipital bone is another
canid feature in common with Bassariscops. This suture lies near the
canal, within one diameter of the canal from the canal itself, in Canis
and Bassariscops, and farther, about two diameters in Procyon and
The great degree of overlap of the auditory bulla, the slitlike medial
terminus of the foramen stylomastoideum definitivum, the grooving of
the mastoid process, the anterior projecton of the petrosal, and the
position of the facial nerve canal are all features shared by Canis and
Bassariscops in contrast to representative mustelids and procyonids.
For these reasons Bassariscops is placed in the Canidae.
The referral of Bassariscops to a particular subfamily of the
Canidae is even more difficult. For reasons outlined below, however, I
am including the genus in the Cynarctinae.
The Subfamily Cynarctinae has accumulated genera rather
haphazardly, and Galbreath (1956) presented a genera review of its
taxonomic history. Hough (1948) first showed Cynarctus, Cynarc-
toides, Phlaocyon, and Aletocyon to be canids without mentioning the
subfamily, and Galbreath (1956), in describing a specimen of Cynarc-
toides acridens in which the basicranial region was typically canid-like,
placed Cynarctoides in the Cynarctinae. In this way Cynarctus,
Cynarctoides, Aletocyon, Phlaocyon, Nothocyon annectans, and now
Bassariscops have become associated with the Subfamily Cynarctinae
within the Canidae. The relationships of these genera are not fully
understood, and they may not all be closely related. At the present
time I think there is merit in having a subfamily of small Miocene
canids which have, as Hough (1948) pointed out, additional cusps and
cingula of all gradations modifying the typical canid carnassial and
molar pattern. Bassariscops, with only a small hypocone on the P4 and
an interoconid as the only accessory cusp on M,, is one of the least
modified genera of the subfamily.

Cynarctoides MCGREW 1938

MATERIAL.-UF 18415, M,.

The M, from Buda resembles that of Cynarctoides acridens
(AMNH 82558, holotype) and C. mustelinus (AMNH 20502, holotype)

Vol. 24, No. 2


in (1) the presence and shape of the opisthoconid (large and clearly
defined to the base of the protoconid in Cynarctoides), (2) the position
of the interoconid (on the crest immediately posterior to the metaconid
in Cynarctoides; in the talonid notch in Nothocyon, except Nothocyon

0 5mm

Xl . .

FIGURE 5.-A) Cynarctoides sp., M,, UF 18415; B) Cynorca sp., M,, UF 18498; C)
Camelidae, n. gen. et sp., P4, UF 18374. Occlusal and labial views of each. Note that B,
Cynorca sp., is drawn to a smaller scale than A and C.





C. acridens C. mustelinus Buda
AMNH 82558 AMNH 20502 UF 18415
length 9.1 8.4 8.4
width of trigonid 4.2 4.0 3.0
width of talonid 4.3 4.0 4.2

annectans); (3) in the greater length of the talonid relative to the
trigonid; and (4) in the large size of the entoconid.
The shape of the opisthoconid is distinctive in Cynarctoides
although the cusp itself is present in other canid groups. It is found on
some specimens of Nothocyon (N. latidens, AMNH 6896, holotype;
6897; and YPM 12794; N. lemur, YPM 12797) but is more generally ab-
sent. Cope (1884) considered this tubercle as diagnostic of N. latidens,
but Thorpe (1922) and Merriam (in Thorpe 1922) found this tubercle on
N. lemur and considered it to be a variable characteristic of the genus.
A slight ridge in the position of the opisthoconid can be seen in
Bassariscops (UF 16989, Fig. 3D), Phlaocyon (AMNH 8768, holotype),
and occasionally in modern canids (Urocyon: Gawne 1973). The occur-
rence of this cusp seems to be within the genetic capability of all
canids, and its presence less indicative of relationship within the fami-
ly than of adaptive response.
The M, from Buda is the most primitive looking tooth referred to
the genus Cynarctoides. It differs from C. acridens and C. mustelinus
in (1) its relatively lower, blunter cusps; (2) in having the entoconid and
hypoconid of nearly equal height (vs. a taller entoconid in the other
species); and (3) in having a wide talonid relative to the trigonid. The
last character is possibly not primitive. This isolated tooth probably
represents a new species of Cynarctoides, but at this time, having as
yet no hypodigm, I prefer to assign this specimen to Cynarctoides sp.


REFERRED MATERIAL.-UF 18409, petrosal; UF 16996, humerus; UF 16927, 3 tibiae;
UF 18404, 2 innominata; UF 18503, femur; UF 18408, calcaneum; UF 18405, metatar-
sal III; UF 18406, 2 metapodials; UF 18407, 12 proximal phalanges.

A large mustelid is represented in the fauna by several
unassociated postcranial elements and one petrosal. Comparisons were

Vol. 24, No. 2


made with postcranial elements of fossil and recent canids, felids, and
mustelids. The size of the postcranial elements is near that of Sthenic-
The petrosal from Buda has features that are uniquely mustelid
among the canoid carnivores, as outlined by Segall (1943). In adult
mustelids the foramen stylomastoideum is almost completely sur-
rounded by the auditory bulla. The tympanic cavity extends well
posteriorly to the promontorium, and there is no indication that the
mastoid portion of the temporal bone forms part of the foramen
Other than the petrosal, the most confidently referrable element is
the humerus (UF 16996). The large blade-like lateral condyloid crest is
exceeded in size only by the unusually large crest on the humerus of
Hoplophoneus; the crest is generally more reduced in carnivore
families other than the Mustelidae. The olecranon fossae of mustelids,
and of the Buda humerus, are intermediate in depth between canids
and felids.
Due to breakage and absence of diagnostic features, the other
elements cannot be definitely assigned to any carnivore family. They
are placed with the mustelid humerus and petrosal because of their
correspondence in size.


REFERRED MATERIAL.-UF 16908, calcaneum.

This Buda calcaneum represents the first Arikareean felid in
Florida. The anteroposterior elongation immediately separates this
calcaneum from the stocky calcanea of canids, procyonids, mustelids,
and machairodont cats. The distance from the lesser process to the
anterior end is equal to one diameter of the articular surface of the
lesser process, and the distance from the posterior process to the
sustentaculum is equal to or greater than the length of the rest of the
calcaneum. The calcaneum from Buda also resembles those of
Nimravinae, but not machairodonts, in its straighter ventral




MATERIAL.-UF 19318, lateral proximal phalanx.

This phalanx is inseparable from those of Parahippus leonensis
from the Thomas Farm Local Fauna but does not constitute sufficient
evidence for a definite taxonomic designation. The extreme rarity of
horses in the Buda fauna is in direct contrast to Thomas Farm, where
horses make up over half the faunal population. This suggests that dif-
ferent environmental parameters were operating at these two localities
during the accumulation of the respective sediments.

GENUS Moropus MARSH 1877

MATERIAL.-UF 24131, M'; UF 16916, M2, ectoloph; UF 24132, trigonid of lower
molar; UF 24133, fragment of lower molar; UF 24129, metatarsal IV; UF 16918,
metatarsal IV, partial; UF 24130, proximal phalanx; UF 16911, medial phalanx; UF
16919, ungual phalanx; UF 16915, ungual phalanx, partial.

The M1 of the Buda chalicothere is remarkable in that it is nearly
quadrangular in outline, a primitive characteristic seen in
Chalicotheriinae and in Schizotherium among Schizotheriinae. Other
schizotherine genera display a quadrangular M' at advanced wear
stages, but UF 24131 has not been shortened by interstitial wear. The
parastyle is large and conical but does not extend anteriorly to the
anterior margin of the tooth, as is also seen in Schizotherium. The pro-
tocone and hypocone have approximately the same height and conical
shape. The protocone and hypocone are connected on their labial edges
by a low crest that divides the central valley. A similar crest occurs in
an unnamed Hemingfordian species of Moropus and rarely if at all in
M. elatus (Coombs 1973:124). Both the protocone and the hypocone
display apical wear. The mesostyle, metastyle, and the paracone are
the tallest cusps on the crown at this wear stage. An external rib is

Vol. 24, No. 2


present on the paracone, and a weak flexure is developed between the
paracone and the parastyle. An anterior valley with a well-developed
anterior cingulum and a postfossette are present. A small lingual
cingulum is present. The length and width of this M' are 21.1 x 20.7.
Although only the ectoloph and metaloph of the M2 are preserved,
its relatively unworn condition permits the description of characters
that are seldom seen on the typically well-worn M2's of Moropus. The
paracone is the tallest cusp on the tooth. The large, conical parastyle is
the next tallest and is intermediate in height between the paracone and
the equally tall mesostyle, metacone, and hypocone. The mesostyle
and metacone are not identifiable as distinct cusps but rather consist
of crests that lie parallel to the occlusal plane before tapering smoothly
to the base of the crown at their outer margins. The metaloph arises
near the mesostyle and quickly drops to a saddle before rising again to
a sharply pointed hypocone. The hypocone is thin and elongate near its
peak but broadens at its base. A small posterior cingulum is present
but only weakly closes the posterior fossette; its greatest expression is
on the posterolabial base of the hypocone. A weak external cingulum is
present labial to the paracone. External ribs are present on the
paracone and on the metacone, although the latter rib is very weak and
would quickly disappear with wear. Shear facets are present on the
lingual surface of the ectoloph anterior and posterior to the paracone,
between the mesostyle and the origin of the metaloph, and on the
lingual side of the metaloph between its origin at the ectoloph and the
hypocone. The length of the ectoloph is 26.0.
The single trigonid of a right lower molar (UF 24132) cannot yield
more than a few features of this species of Moropus. In most respects,
this trigonid is similar to those of M. elatus and may be the trigonid of
an M2, judging by the prominence of the paraconid, the V-shaped
valley of the trigonid basin, and the relatively large, symmetrical crest
that closes the lingual border of the trigonid basin. The length (antero-
posteriorly) of the trigonid is 10.3; the width is 13.0.
Two metatarsals IV (UF 16918, 24129) are preserved in the Buda
sample. UF 16918 lacks the proximal end. These two metapodials are
unequal in size and evidently represent two size groups in the popula-
tion, a situation known to occur among chalicotheres (Coombs 1975).
The proportions of the metarsal IV are very similar to those of
Moropus elatus. The articular facets for metatarsal III are separate,
with the volar facet about one-half the size of the dorsal facet. There is
no articular facet for the ectocuneiform on the dorsal facet.
The cuboid articulation is rectangular and separated from the
facets for metatarsal III by a sharp crest. The fibular-volar pro-
tuberance is well developed as in M. elatus. The shaft is square in




0 2cm

0 2cm

Vol. 24, No. 2



mu 4v Y


FIGURE 6.-Moropus sp. A-F: left metatarsal IV, UF 24129. A) dorsal view; B) medial;
C) volar; D) lateral; E) proximal end; F) distal end. G-L: left proximal phalanx, UF
24130. G) dorsal view ; H) medial; I) volar; J) lateral; K) proximal end; L) distal end.
M-R: right medial phalanx, UF 16911. M) dorsal view; N) lateral; 0) volar; P) medial; Q)
proximal end; R) distal end. S-X: left ungual phalanx, UF 16919. S) dorsal view; T)
medial; U) volar; V) lateral; W) proximal end; X) distal end. Y) partial ungual phalanx,
UF 16915, proximal end. AA) left M', UF 24131, labial view; BB) UF 24131, occlusal
view. CC) partial left M2. UF 16916, labial view; DD) UF 16916, occlusal view. Large
scale refers to teeth only.

cross-section proximally, whereas in specimens of M. elatus the shaft
is roughly triangular for the distal four-fifths of its length.
Measurements of metatarsals IV, (UF 24129, 16918), are as
follows: maximum length 93.7; proximal end, depth 24.1, width 30.0;
distal end, depth 28.5, 23.2 (UF 16918), width, 26.2, 21.0 (UF 16918);
mid-shaft, minimum depth 17.9, 13.0 (UF 16918), minimum width
18.0, 15.4 (UF 16918); length/distal width ratio 3.6; length/shaft width
ratio 5.2.
The proximal phalanx (UF 24130) closely resembles the left prox-
imal phalanx of pedal digit III of Moropus elatus illustrated by
Holland and Peterson (1914:375). Measurements of the proximal
phalanx are: length 45.7; proximal width 26.5; distal width 18.5; depth
at dorsal margin of metapodial facet 21.3; depth at volar termination
of distal articular surfaces 16.8.
The medial phalanx (UF 16911) displays a slight asymmetry in the
proximal articular surfaces in that one facet, the medial?, is longer
than the other. The distal articular surfaces are symmetrical. The size
of this medial phalanx is surprisingly large, approaching that of M.
elatus females (=M. "petersoni"; Coombs 1975). Measurements are as
follows: length 28.1; width of distal articular surface at the dorsal ter-
mination, 13.2; at the volar termination 17.6; dorsal to volar depth of
the distal end 23.9.
The proximal end of one ungual phalanx (UF 16915) is flatter and
more asymmetrical than another (UF 16919) and apparently is from a
marginal digit. The more complete ungual phalanx (UF 16919), miss-
ing only a portion of the dorsal surface, cannot confidently be referred
to a particular digit of the pes or manus. This ungual does not narrow
proximal to the cleft and has a large dorsal process. The claw sides of
the phalanx are deeply rugose and, in volar view, widely separated (6
mm). Measurements of the ungual phalanges are: maximum length
51.3; greatest width 20.7; height 24.5; volar depth of cleft 14.7; dorsal
depth of cleft 27.3; height and width of articular surfaces 18.6 x 16.2
(UF 16919), 10.5 x 17.2 (UF 16915).



In the quadrangular M', and probably M2 and M' as well, and the
absence of an ectocuneiform contact on the metatarsal IV, the Buda
chalicothere resembles Schizotherium, a primitive Eurasian Oligocene
schizotherine (Coombs 1973). These features, however, are primitive in
the Chalicotheriidae and hence of dubious taxonomic importance at
this level. The metatarsals of Schizotherium, as seen in S. priscum and
S. turgaicum (Coombs 1974, 1978) are far more slender than those of
the Buda chalicothere.
In the lack of an ectocuneiform facet on the metatarsal IV and in
the combined convexity of the facets for metatarsal III, the Buda
chalicothere is similar to an unnamed new genus of North American
Schizotheriinae (Coombs, in press) in which the metapodials are much
more shortened and stocky, but in which the upper molars are more
strongly elongated.
Small species of Moropus are known from earliest Miocene deposits
of North America and Europe, although available material is fragmen-
tary. Moropus oregonensis, from the John Day Basin of Oregon (Leidy
1873) is known only from teeth: Moropus distans, also from the John
Day (Marsh 1877), is known only by podial elements; and Moropus sp.
from St. G6rand-le-Puy, France, is known only from a partial ramus of
a juvenile and a metatarsal IV (Coombs 1974, in press). The Buda
material cannot be compared with that of M. distans, and it is neither
M. oregonensis nor Moropus sp. from St. Grand. It differs from M.
oregonensis in having a quadrangular M' and a larger, more labially
placed mesostyle on M'. The metatarsal IV of the Buda chalicothere
differs from that of Moropus sp. from St. Grand in that it lacks an ar-
ticular facet for the ectocuneiform, and it has a smaller and more con-
vex volar facet for metatarsal III. Coombs (in press) suggests that M.
distans may be conspecific with M. oregonensis on the basis of their
small size and sympatry. If this proves to be the case, then I doubt
that the Buda chalicothere can be referred to this species because of
differences already noted between the teeth of M. oregonensis and the
Buda chalicothere.
The Buda chalicothere is apparently a primitive schizotherine
chalicothere in that it does not have strongly elongated upper molars
and the length and sturdiness of the metatarsals is relatively un-
modified. The metatarsals IV are most similar to those of Moropus
elatus and for this reason the generic identification is given to

Vol. 24, No. 2


Cynorca COPE 1867

MATERIAL.-UF 18498, M,; UF 18496, fragment of innominate; UF 18495, frag-
ment of metapodial; UF 19317, 5 proximal phalanges; UF 18497, 2 medial phalanges.

The simplicity of the cusps and the small size of the M, from Buda
(Fig. 5B) are characteristic of Cynorca, the smallest of the Oligocene-
Miocene peccaries. The M, from Buda is more elongate than that of C.
social and the cusps are less swollen. In these features the M, from
Buda is like that of C. proterva. However, the diagnostic post-
metaconid ridge of C. proterva is not evident on the Buda M, and the
M, of C. proterva appears wider. The narrowness of the Buda M, may
be due partly to breakage at the base of the crown. The length and
width of the M, from Buda is 11.8 x 6.8.

The only other Arikareean occurrences of Cynorca presently known
are C. social from the John Day fauna of Oregon, and a single
specimen, Cynorca cf. social from the "Loup Fork Tertiary" of
Nebraska (Woodburne 1969)'. The latter may in fact be later than the
Cynorca proterva is known from the Barstovian of Maryland,
Texas, and Nebraska, with a questionable late Hemingfordian occur-
rence in Nebraska (Woodburne 1969).
Woodburne (1969) stated that Cynorca proterva is primarily an
eastern and Gulf coastal species that probably arose from C. social in
the early or middle Miocene. The single molar of Cynorca from Buda,
which bears features of both C. social and C. proterva, supports this

'Woodburne (1969) stated that there are few stratigraphic data associated with this specimen. The Loup Fork is an
obsolete term that was once in general use for part or all of the Miocene (Simpson 1933).


0 2cm C

FIGURE 7.-Merycoidodontidae, Phenacocoelinae gen. et sp. indet. A) Right P", UF
18424; B) Left P', UF 16903; C) Right M3, UF 16931. Occlusal views.


REFERRED MATERIAL.-UF 18423, C,; UF 16902, C'; UF 18424, P3; UF 16903, 2 P4's;
UF 16932, M'; UF 16931, 2 M3's; UF 18428, 2 incisors; UF 16979, teeth fragments; UF
19319, auditory bulla; UF 18336, petrosal; UF 17007, axis vertebra; and UF 18420,
18421, 18425-18427, 18430, 18432, various postcranial elements.

Unlike most fossil mammals, the taxonomy of the oreodonts is
largely based on cranial anatomy and not on dental features. Because
of the critical lack of cranial materials from Buda, precise identifica-
tion of this oreodent is not possible'. Some preliminary statements,
however, can be made from the dentition alone, especially from the P3.
Although there is a great amount of individual variation among
oreodonts in the shape of P3, members of the Phenacocoelinae, in con-
trast to other subfamilies of oreodonts, have an anteroposteriorly com-
pressed P3, which approaches P4 in outline. The anteroposterior com-
pression of P3 is evident in the reduction, almost to insignificance, of
the anterior crescent of this tooth. The flexure of the labial crest of the
P3 from Buda (Fig. 7A) is more concave than is commonly seen in any
subfamily of oreodonts, including the Phenacocoelinae.
Within the Phenacocoelinae, the nearest comparison of the Buda
material is with Phenacocoelus stouti (Schultz and Falkenbach 1950).
The P", M', and M3 of P. stouti are inseparable from those of the Buda

'Dr. Bruce Lander, in the course of his studies of oreodonts at the University of California, examined the oreodont
material from Buda. Although he concurred with the identification of the dental material, he felt the isolated auditory
bulla (UF 19319) was more like that of Merycochoerus matthewi from the lower part of the Marsland Formation of

Vol. 24, No. 2


oreodont. The P3, although close, does not have the same degree of
anteroposterior compression as is seen in the Buda P3. Also, the
anterior crescent of P3 is slightly smaller and the exterior crescent is
more concave on the P3 from Buda than on that of P. stouti. A new
species of Hypsiops from the Big Bend area of Texas may also be
related to the Gulf Coastal phenacocoelines (Stevens et al. 1969).
The Subfamily Phenacocoelinae contains four genera: Phenacocoe-
lus, Hypsiops, Submerycochoerus, and Pseudomesoredon. All species
of the four genera, except Phenacocoelus stouti, are strictly
Arikareean species. Phenacocoelus stouti is a Hemingfordian species.
The subfamily is widely distributed over the western part of the
United States (Schultz and Falkenbach 1950), but was unknown in the
southern states until Stevens et al. (1969) described a new species,
Hypsiops leptoscelos, from the Castolon Local Fauna (Arikareean) of
the Big Bend area of Texas. The oreodont from Buda is the first occur-
rence of the Phenacocoelinae in the Gulf Coastal Plain.


REFERRED MATERIAL.-UF 18373, 6 incisors; UF 17015, 8 incisors; UF 18382, 10
premolars; UF 18385, 3 P4's; UF 18384, 12 upper molars; UF 18378, 25 upper molar
fragments; UF 19313, 2 P,'s; UF 18365, 3 P,'s; UF 18387, 3 P4's; UF 18374, P, with
mandibular fragment; UF 18379, 19 lower molar fragments; UF 18388, 7 lower molars;
UF 19314, 2 M,'s; UF 22779, 4 DP3's; UF 22780, 3 DP3's; UF 19315, 6 deciduous lower
premolars; UF 16950, 7 deciduous teeth; UF 18384, 4 mandibular fragments; UF 16907,
16914, 16955, 17006, 18367-18369, 18370-18372, 18375-18377, 18380, 18381, various
postcranial elements.

COMPARATIVE MATERIAL ExAMINED.-Floridatragulus dolichanthereus, F:AM
31864, 31865; Gentilicamelus sternbergi, AMNH 7910, holotype; Miolabis transmon-
tanus, AMNH 8196, holotype; Miolabis sp., F:AM 68985, 68986; Miotylopus
bathygnathus, MCZ 2924, holotype (cast in AMNH); Nothokemas floridanus, UF
19929, AMNH 22672; Oxydactylus campestris, AMNH 17620, holotype; Oxydactylus
longipes, CM 918, holotype (cast in AMNH); Paratylopus primaevus, AMNH 9806,
holotype; Poebrotherium eximium, AMNH 632, holotype.

Excluding the oromerycids, this is the smallest known camel, equal
in size to Leptomeryx or Hyemoschus. These specimens probably
represent a new genus of camel. I am, however, reluctant to erect a new



genus on the basis of isolated teeth, because the possibility exists that
future researchers may recognize another taxon in this sample or not
be able to differentiate species of a genus whose holotype is an isolated
Among the Camelidae, this small camel from Buda may be related
to a group comprised of Oxydactylus, Nothokemas, Gentilicamelus,
and perhaps Floridatragulus and Miolabis. Each has characteristic
features of its own, but all share a distinctive enamel pattern in the
molars, especially evident in the lower molars, in contrast to all other
camelid genera.
UPPER DENTITION.-The DP3's from Buda resemble those of Ox-
ydactylus but differ from those of Miolabis, which are shortened and
have only one prominent cusp not two.
P" of the Buda camel is very similar to P4 of all camelids.
The upper molars of the Buda camel (Fig. 8H, I, K, L) also resemble
those of Oxydactylus, Gentilicamelus, Miolabis, Nothokemas, and
Floridatragulus. In appearance, the metacone (posterior crest) slightly
overlaps the paracone (anterior crest) with the forward edge of the
metacone turning out sharply to form a prominent mesostyle. With
wear the crests unite first and then the crescents. The upper molars of
each genus have prominent mesostyles and external ribs. Small inter-
columnar styles are present on molars of the Buda camel, Gen-
tilicamelus, and Floridatragulus and are individually variable in size
and presence in Oxydactylus and Miolabis.
LOWER DENTITION.-P2 and P, (Fig. 8A, B) have simple enamel pat-
terns lacking lingual stylids in contrast to those of Gentilicamelus and
Nothokemas. A lingual stylid may or may not occur on P, of Oxydac-
tylus. A DP3 found at Buda (Fig. 8F) has a prominent, curved lingual
stylid as does DP, of Oxydactylus. DP3 of Miolabis has a straight
lingual stylid. Deciduous lower premolars of Nothokemas,
Floridatragulus, and Gentilicamelus are unknown.
P4 of the Buda camel (Figs. 5C, 8C) is diagnostic. This tooth is not
wedge-shaped as are P4's of most other camelid genera. The width of P4
at the protoconid is approximately equal to the width at the hypoconid
giving the tooth a quadrate outline. This is also seen in
Floridatragulus and in some individuals of Miolabis.
The enamel pattern of the lower molars is shared by this new genus
(Fig. 8D, E, G) and Oxydactylus, Gentilicamelus, Nothokemas,
Floridatragulus, and Miolabis. The crests of the lower molars are not
in the same vertical plane but rather in parallel planes, the metaconid
(anterior crest) slightly overlapping the entoconid (posterior crest).
The metaconid remains free of the entoconid, the hypoconid (posterior
crescent), and the posterior end of the protoconid (anterior crescent)

Vol. 24, No. 2

0 5mm /1 ,


0 2cm

V __iK L

FIGURE 8.- Camelidae n. gen. et sp. A-E: Composite left lower tooth row. A) P,, UF 19313; B) P,, UF 18365; C) P,, UF 18387; D) Lower
molars, M, reversed, UF 18385; E) Composite M3 using M3, UF 19314, and lower molar, UF 18385; F) Left DPs,UF 22780; G) Left M, or M2,
UF 18388; H, I, K, L) Upper molars, UF 18384, (H-I. Labial views, K and L, Occlusal views, H and K are reversed); J) Left metacarpal III,
UF 18367, natural size.


until advanced wear of the tooth. The protoconid and entoconid are the
first parts of the tooth to unite after wear. Slight internal ribs are pres-
ent on the lower molars of all six genera. Intercolumnar styles are pre-
sent on Gentilicamelus, the Buda camel, and Nothokemas and, as in
the upper molars, variably present in Oxydactylus and Miolabis.
MANDIBLE.-The lower ramus of this new genus has a longer
diastema between P, and P2 than does Gentilicamelus sternbergi. The
ramus is sharply constricted in this area as in Oxydactylus.
POSTCRANIAL SKELETON.-The postcranial skeleton of the Buda
camel is difficult to separate from skeletal elements of other
Arikareean camels on features other than size. The metapodials are un-
fused and of the same proportion as those of Gentilicamelus. A single
complete metacarpal III (UF 18367; Fig. 8J) measures 94.7 mm in
length. None of the elongation of neck and limbs typical of Oxydac-
tylus is seen in the skeleton of the new genus from Buda.

During the Arikareean and the Hemingfordian, the generic diver-
sity of camels with low crowned teeth was greater in the Gulf Coast
than anywhere else in North America. In the Arikareean, ?Oxydac-
tylus or ?Nothokemas (Simpson 1930) and this new genus from Buda
were present in Florida. The second camel found at Buda, discussed in
the next section of this paper, may be yet another genus. Further
discussion of these camels is deferred to that section.


REFERRED MATERIAL.-UF 19316, upper molar; UF 18386, 2 lower molars; UF
18364, 3 fragments of scapulae; UF 18362, 5 fragments of humeri; UF 18358, 5
fragments of radii; UF 18363, 2 fragments of innominata; UF 18361, 3 fragments of
sacra; UF 18359, 6 fragments of femora; UF 18360, 6 fragments of tibiae; UF 16956, 10
astragali; UF 18383, 2 fragments of metapodials; UF 16925, 53 metapodials; UF 18366,
70 phalanges.

A second camel, about twice the size of the camel discussed in the
previous section, is represented in the Buda Local Fauna by numerous
fragmentary postcranial elements. Only two complete teeth, an upper
molar and a lower molar, can be referred to this taxon. Each tooth,
however, has the pronounced overlapping parallel crests that are seen

Vol. 24, No. 2


0 5mm


FIGURE 9.- Camelidae, gen. et sp. indet. A) Right upper molar, UF 19316, occlusal and
labial views. B) Left lower molar, UF 18386, occlusal and lingual views.

in Oxydactylus and a few other genera, as is discussed in the previous
section and again below.
The upper molar is an M2. The degree of brachydonty and overlap
of the paracone by the metacone is comparable to that seen in Gen-
tilicamelus and Floridatragulus. A small intercolumnar style is pres-
ent and resembles those of Floridatragulus, the unnamed new genus
from Buda, and Gentilicamelus. The external ribs and styles, however,
are less pronounced than in Floridatragulus or Gentilicamelus, but
resemble those in Nothokemas.
The lower molar (Fig. 9B) lacks an intercolumnar cingulum. In the
great degree of overlap of the entoconid (posterior crest) by the
metaconid (anterior crest), this camel is very much like the unnamed
new genus from Buda. The internal ribs of the lower molars are more
pronounced than in any other camelid genus examined, including Gen-
tilicamelus and Floridatragulus.
The proximal ends of broken metapodials suggest that the




metapodials were not fused. This is a primitive tylopod feature also
observed in Oxydactylus and Gentilicamelus.

The low crowned teeth with an Oxydactylus-like pattern can be
compared with Oxydactylus, Gentilicamelus, Nothokemas, Miolabis,
Floridatragulus, and the unnamed new genus previously described. All
occur in the Gulf Coastal Plain, and Floridatragulus, Nothokemas, and
the new genus are restricted to the Gulf Coast.
The two camels from Buda both have brachydont molars and un-
fused metapodials. The retention of these primitive camelid features in
both species suggests that selection pressures on these camelid
populations in Florida during the Arikareean differed from those on
many camelids in western North America where hypsodont molars
and fused metapodials were more common. Presumably Buda sampled
a forested region where there was no great emphasis on a harsh diet
and sustained running.

Nanotragulus LULL 1922
Nanotragulus loomisi LULL 1922

REFERRED MATERIAL.-UF 16953, 29 incisors; UF 16937, 25 P2's; UF 18482, 14 P"'s;
UF 16985 14 P^'s; UF 16994, 62 upper molars; UF 18418, M2 embedded in limestone;
UF 18500, M'-3; UF 17014, 14 M's; UF 18483, 38 P,'s; UF 18484, 25 P,'s and P,'s; UF
16960, 19 P,'s; UF 16958, 43 lower molars; UF 18485, 15 M,'s; UF 18486, 2 fragments of
maxillae; UF 16959, 19 fragments of mandibles; UF 16913, 16924, 16949, 16951, 16997,
16998, 18487-18494, various postcranial elements.
COMPARATIVE MATERIAL EXAMINED.-Nanotragulus loomisi: YPM 10330, holotype.
N. cf. loomisi: SDSM 5995, 53397, 53402, 53432, 54340, 59109. N. "lulli": AMNH
13821, holotype. N. "intermedius": MCZ 2301, holotype; MCZ 2812, paratype; and
numerous referred specimens at SDSM from the Sharps Formation of South Dakota.
Three genera of hypertragulids and one leptomerycid (sensu Taylor
and Webb 1976) have unfused metapodials and approximate the size of
the material from Buda. These are Nanotragulus, Hypertragulus,
Hypisodus, and Leptomeryx. Each genus has distinctive dental
features which permit certain identification, even though only a few
isolated teeth may be available (see Table 5). Comparisons of these
four genera are presented in this section to illustrate the differences

Vol. 24, No. 2

TABLE 5.-DENTAL FEATURES THAT DIFFERENTIATE Nanotragulus, Hypisodus, Hypertragulus, AND Leptomeryx.

Nanotragulus Hypisodus Hypertragulus Leptomeryx

P': lingual fossette, after wear 2, almost equal in size 2, equal in size 2, posterior smaller 1
Crown height: subhypsodont subhypsodont brachydont brachydont
Upper molars: mesostyles absent absent minute present
intercolumnar styles small, often absent small, often absent minute, small, variable present, small, variable
intercolumnar cingulum absent absent present present but very small
M3 metastyle large, large, small, small,
leans anteriorly leans anteriorly parallel to external rib parallel to external rib
P,: metaconid indistinct distinct distinct distinct
Lower molars:
intercolumnar stylids variable in size
and presence usually absent small absent
intercolumnar cingula absent usually absent present small, usually absent
M3 posterior fossettid closed closed closed open
Size: relative to Nanotragulus 1.0 0.7 1.0 1.2


among them and the reasons for the subsequent referral of the Buda
material to Nanotragulus.
UPPER DENTITION.-Although variable, the characteristics of the
P4 of all four genera listed above are roughly similar. The protocone of
Nanotragulus is displaced anteriorly on the P'. P4 of Nanotragulus has
a small tubercle which, with a little wear, becomes a ridge connecting
the protocone medially with the ectoloph and dividing the central
fossette of an unworn tooth into two slightly unequal fossettes (the
anterior being larger) that persist until advanced wear. P4 of Hyper-
tragulus is like that of Nanotragulus, except that the two fossettes in
Hypertragulus are very unequal in size, the smaller posterior fossette
quickly disappearing with wear. In contrast to Nanotragulus, P4 of
Leptomeryx has a more medially located protocone and has only one
fossette, never two, although the posterior part of the single fossette
wears away first, giving the appearance of there having been at one
time two fossettes. P4 of Hypisodus, unlike Nanotragulus, has a large,
medially placed protocone that after slight wear is connected to the ec-
toloph by a crest that divides the single fossette of an unworn tooth in-
to two nearly equal fossettes that persist until advanced wear.
The upper molars of Nanotragulus and Hypisodus are equally
subhypsodont and lack mesostyles and internal cingula. An inter-
columnar style is variable in both presence and size. In contrast, the
upper molars of Leptomeryx are brachydont with large parastyles,
mesostyles, and metastyles. A small intercolumnar style rises from an
intercolumnar cingulum on the metaconule of each upper molar of Lep-
tomeryx. These styles and cingula vary in size but are never large or
prominent. The upper molars of Hypertragulus are also brachydont,
have incomplete internal cingula (larger than those seen on the upper
molars of Leptomeryx, yet they do not extend over the most lingual
surface of each cusp), and there is a minute mesostyle. The presence
of intercolumnar cingula and mesostyles on the upper molars of
Hypertragulus contrast this genus with Nanotragulus. Lull (1922) and
Cook (1934) stated that the upper molars of Hypertragulus have no
mesostyles, but in fact minute mesostyles are present.
In M3 of Hypertragulus the metastyle is parallel to the external
ribs for most of its length, and Leptomeryx has a similar small
metastyle. In contrast, the metastyles of Nanotragulus and Hypi-
sodus are large and lean anteriorly (the teeth widen towards their
bases) instead of being parallel to the external ribs.
LOWER DENTITION.-P4 of Nanotragulus is extremely variable (Fig.
10E) but most similar to that of Leptomeryx. The metaconid is never
separate as in Hypertragulus and Hypisodus; instead it is difficult to
differentiate from the inner enamel wall of the P4.

Vol. 24, No. 2


0 5mm

FIGURE 10.-Nanotragulus loomisi. A) Fragment of left mandibular ramus with M, 2,
UF 16959, occlusal view; B,D) Left M,, UF 18485, occlusal and labial views; C) Two
fragments of left mandibular rami, UF 16959, labial view, showing variation in the P,-P,
diastema, lower specimen with P,-P4; E) Five left P,'s, UF 16960, showing variation in
enamel pattern; F) Left M2'-, UF 18500, occlusal view.

The lower molars of Hypertragulus have intercolumnar cingula. On
Ms the cingulum is variable in extent and may appear more as a style
between the hypoconid and the hypoconulid. It is usually absent on
the posterior part of the hypoconulid. The intercolumnar labial cingula
of Nanotragulus are vertically accentuated into thin styles (Fig. 10D).
These are variable both in size and presence. The intercolumnar
cingula of Hypisodus are very reduced and usually absent altogether.
The cingula and styles on the lower molars of Leptomeryx are variable


in size yet smaller than in Hypertragulus or Nanotragulus. Few lower
molars of Leptomeryx have intercolumnar cingula and none have in-
tercolumnar styles. Leptomeryx differs from the hypertragulids com-
pared in having an open posterior fossette on M3.

The specimens from Buda fall within Nanotragulus Size Group IV'
of Frick (1937). Size is certainly the easiest and may even be the most
reliable way to identify the species of Nanotragulus, but other dif-
ferences between N. loomisi and species in other size groups are
Nanotragulus albanensis Frick (1937) (F:AM 13785, holotype) is 50
percent larger than the Buda species. The lower molars have larger in-
tercolumnar tubercles than those of N. loomisi (especially M3, which
has two).
Nanotragulus ordinatus (Matthew 1907) (F:AM 13013, 13011) is 25
percent larger than the Buda species. P, of all N. ordinatus specimens
examined differ from the Buda specimens in having the major cusp
centrally placed between the roots and not over the anterior root. The
inner selenes of the lower molars, especially those of N. ordinatus var.
(sensu Frick 1937) (F:AM 31534-6), are more rounded than those of N.
Nanotragulus matthewi Cook (1934) compares in size to N. or-
dinatus (Stevens et al. 1969).
Frick (1937) included two species, N. loomisi Lull (1922) and N. lulli
Frick (1937) in his Size Group IV, the smallest size group. Schlaikjer
(1935) named a new species, N. intermedius, which was briefly men-
tioned but not discussed by Frick (1937) and which also falls in this
size group. The amount of variation seen in the teeth and postcranial
elements of Nanotragulus material from Buda, a very restricted sam-
ple both temporally and geographically, casts doubt on the diagnostic
charactersitics used by Frick (1937) and Schlaikjer (1935) to
distinguish these two species from N. loomisi.
While Frick (1937) diagnosed N. lulli on the basis of the large
auditory bulla, the longer limb elements, and the slightly larger size of
the type specimen, the teeth are morphologically inseparable from
those of N. loomisi. Frick (1937:643) stated that the bulla of N. lulli is
large in comparison to Hypertragulus. This is true, however, for all
species of Nanotragulus and is not diagnostic of a single species. A
tibia from Buda (UF 18493) measures 92 mm and is thus closer to N.
loomisi. On the other hand, the third metatarsal of N. loomisi
'Frick (1937) grouped species by size as a convenient first step in systematic discussion. The number of groups or the
size range within a group differed with each major taxon.

Vol. 24, No. 2


measures 50 mm; that of N. lulli measures 56 mm. Two third metatar-
sals from Buda both measure 50 mm, but a metatarsal IV (which is
usually within 1 mm of the matching metatarsal III) is 54 mm, bridg-
ing the gap in metatarsal measurements between the two species. The
lengths of the limb elements of Nanotragulus from Buda do not cluster
around comparable measurements of either N. loomisi or N. lulli, nor
are they consistently larger or smaller than these two species. The
variation in limb lengths lends support to the contention that this is
due to individual variation and is not diagnostic of a single species of
Nanotragulus. Other measurements of the postcranial elements of
Nanotragulus from Buda indicate a wide size range in this population,
overlapping samples referred by Frick (1937) to N. loomisi and
samples of N. lulli (Table 6).
Much of Schlaikjer's (1935) diagnosis of N. intermedius was made
on cranial elements and complete tooth rows, which cannot be com-
pared for lack of counterparts in the Buda specimens. The diagnostic
characteristics that pertain to teeth and size, however, are all variable
within the limits of the Buda Nanotragulus material. Dental
measurements of Nanotragulus loomisi, N. "lulli" (=N. loomisi), and
N. intermedius (Table 7) do not differentiate these species, while coeffi-
cients of variation between 4 and 10 indicate a unified sample (Simp-
son et al. 1960). The consistency of the coefficients of variation of
Nanotragulus is remarkable since these specimens represent deposits
of similar but not identical ages from localities in Florida, Wyoming,
and South Dakota. A scatter diagram of upper and lower third molars,
the largest sample of recognizable isolated teeth from Buda (Fig. 11),

N. loomisi (=N. lulli; in Loomis 1933) FROM PORCUPINE CREEK, SOUTH

Spanish Diggings Porcupine Creek Buda
Greatest internal
diameter of
acetabulum 7.4 8.9 10.0 N=2
Greatest length
of tibia 106.2 118 92.4
Maximum width
of distal end
of humerus (9.9) 10.0- 11.4 N=6
Length of
metatarsals (50.3 50.9) N=2 56 50.0- 54.4 N=3
Length of
astragali 10.4 10.9 N=2 10.8 12.5 N=14



M' (length)
N. loomisi (YPM 10330, holotype)
N. "lulli" (AMNH 13821, holotype)
N. "intermedius" (MCZ 2103, holotype)
N. "intermedius" (MCZ 2812, paratype)
N. "intermedius" (Sharps Fm.)
N. cf. loomisi (Sharps Fm.)

N. loomisi (YPM 10330, holotype)
N. "lulli" (AMNH 13821, holotype)
N. "intermedius" (MCZ 2103, holotype)
N. "intermedius" (MCZ 2812, paratype)
N. "intermedius" (Sharps Fm.)
N. cf. loomisi (Sharps Fm.)
M3 (length and width)
N. loomisi (YPM 10330, holotype)

N. "lulli" (AMNH 13821, holotype)

N. "intermedius" (MCZ 2103, holotype)

N. "intermedius" (MCZ 2812, paratype)


1 11.3
1 13.3
1 12.5
1 12.4
7 11.3-13.9
3 11.3-12.1
14 11.3-13.9

L 1
L 1
L 1
L 1






N. loomisi (Buda) L 15 6.7-8.1 7.3 0.381 5.229
W 5.2-6.3 5.9 0.376 6.396
N. "intermedius" (Sharps Fm.) L 7 6.3-7.2 6.8 0.298 4.406
W 5.1-6.5 5.8 0.535 9.231
N. cf. loomisi (Sharps Fm.) L 2 6.1-6.7 6.4 0.424 6.629
W 5.7-5.8 5.8 0.070 1.229
all (except MCZ 2103) L 26 6.1-8.1 7.1 0.454 6.416
W 4.8-6.5 5.8 0.443 7.655
All (incl. MCZ 2103) L 27 6.1-8.1 7.1 0.464 6.579
W 4.4-6.5 5.7 0.507 8.850

M, (length and width) t
N. loomisi (YPM 10330, holotype) L 1 4.8 0
W 3.2 >
N. "lulli" (AMNH 13821, holotype) L 1 4.8
W 3.9
N. "intermedius" (MCZ 2103, holotype) L 1 5.0
W 3.3
N. loomisi (Buda) L 1 5.1
W 3.5
N. "intermedius" (Sharps Fm.) L 14 3.9-4.9 4.4 0.338 7.704
W 3.2-3.9 3.4 0.216 6.315
N. cf. loomisi (Sharps Fm.) L 2 4.6-4.8 4.7 0.141 3.008
W 3.3-3.6 3.4 0.212 6.148
all L 20 3.9-5.1 4.5 0.364 8.042
W 3.2-3.9 3.4 0.223 6.489


M,-3 (length)
N. loomisi (YPM 10330, holotype)
N. "lulli" (AMNH 13821, holotype)
N. "intermedius" (MCZ 2103, holotype)
N. "intermedius" (Sharps Fm.)
N. cf. loomisi (Sharps Fm.)
all (except MCZ 2103)
all (incl. MCZ 2103)

M3 (length and width)
N. loomisi (YPM 10330, holotype)

N. "lulli" (AMNH 13821, holotype)

N. "intermedius" (MCZ 2103, holotype)


L 1
L 1
L 1
L 5
L 15
L 3
L 25
L 26

N. loomisi (Buda)

N. "intermedius" (Sharps Fm.)

N. cf. loomisi (Sharps Fm.)

all (except MCZ 2103)

all (incl. MCZ 2103)








7.035 .


also fails to separate Nanotragulus intermedius from N. loomisi and N.
"lulli" (=N. loomisi).
Other diagnostic features used by Schlaikjer (1935) are equally in-


oLn A
00 n N


5.0 [

Sholotype (approx.)

6.0 .5 7.0 .5
Length M3

0 -

o c oo0
001 OD

3.0 [

Sholotype (approx.)

.5 7.0 .5 8.0 .5
Length M3

9.0 .5

FIGURE 11.-Scatter diagrams of M3 and M3 of Nanotragulus. Squares = N. loomisi;
circles = N. "intermedius"; hexagons = N. "lulli"; triangles = Nanotragulus from
Buda. Solid symbols represent holotypes or paratypes. Measurements are in

8.0 .5

4.0 [



conclusive. The upper molars vary greatly in outline. They may be
quadrangular (as Schlaikjer [1935] described the molars of N. in-
termedius), or the metacone may be more labially displaced giving an
uneven appearance to the tooth. The internal cusps of the P's from
Buda are variable in size. Some have large inner cusps while in others
(5 of 14) this cusp is reduced to a cingulum. Schlaikjer (1935) described
the P3 of N. intermedius as having a small internal cusp. In the
holotype of N. intermedius (an immature individual in which M3 and
Ms have not fully erupted) the M3 does not appear to overlap M2 as
much as in specimens of N. loomisi, but in the paratype of N. in-
termedius (MCZ 2812, an adult), the degree of overlap is as great as
that seen in comparable specimens of N. loomisi. The degree of overlap
of M2 by M3 appears to be highly variable within the Buda population
of Nanotragulus. Is is not larger than I, or I2 on N. intermedius as
Schlaiker (1935) stated in the diagnosis, but is smaller as Schlaikjer
noted in the description. In either event the difference is slight and
variable. In N. intermedius, P2 and P, are said to be closely appressed.
In the Buda sample, four mandibular fragments have no diastema be-
tween P2 and P, and two others have small diastemata, the largest
equalling P2 in length (Fig. 10C). The presence or absence of a diastema
between P2 and P, seems also to be a variable feature in Nanotragulus
populations. The presence or absence of intercolumnar pillars is
another variable feature of the teeth of Nanotragulus. Lull (1922)
stated that they were absent in N. loomisi. Schlaikjer (1935) found
them on the holotype of N. intermedius but absent on the paratype. Of
60 complete upper molars of Nanotragulus from Buda, 6 (one M3) have
definite intercolumnar pillars. Of 19 complete lower molars (no M,'s)
from Buda, 5 have a small intercolumnar pillar.
The small size attributed to N. intermedius in the sample examined
by Schlaikjer (1935) appears to be due to one small individual (the
paratype, MCZ 2812), and an immature individual (the holotype, MCZ
2013). From those measurements and morphological characters that
could be compared between the holotype and paratype of N. in-
termedius, the holotype of N. loomisi (YPM 10330), referred specimens
from South Dakota, and the sample from Buda, it seems that only one
species can be recognized. By priority that species is Nanotragulus
loomisi Lull 1922.


The age of the Buda Local Fauna is difficult to place in that of four
species that could be accurately identified, three are new. Because

Vol. 24, No. 2


Nanotragulus loomisi has been recognized elsewhere in the United
States, it is the best age indicator in the fauna.
The holotype of Nanotragulus loomisi was described from Castle
Butte, near Spanish Mines, Wyoming (Lull 1922). Lull (1922) con-
sidered these beds to be "Lower Harrison" equivalents. Loomis (1933)
considered Lull's "Lower Harrison" beds as equivalent to "Lower
Rosebud" beds of South Dakota. Schlaikjer (1935) went even further
and regarded Lull's (1922) Spanish Mines "Lower Harrison" and the
"Lower Rosebud" as facies of the same rock unit. These "Lower
Rosebud" beds of South Dakota are now known to include beds from
the upper part of the Sharps Formation through the lower part of the
Harrison Formation (Macdonald 1963). Macdonald (1963) found
Nanotragulus loomisi in the Sharps Formation only. According to
Macdonald (1963), the Sharps Formation is the lower-most Arikareean
stratum in the Wounded Knee area and is overlain by the Monroe
Creek and Harrison formations. Although other Nanotragulus species
may occur in the Whitneyan and Hemingfordian deposits, the genus is
most frequently found in deposits of Arikareean age (Frick 1937). The
presence of N. loomisi in the Buda Local Fauna gives the fauna a
decidedly Arikareean aspect.
The presence of Daphoenodon notionastes and its stage of evolu-
tion also indicate a late Arikareean or, at the latest, a very early Hem-
ingfordian age for the Buda Local Fauna. It is not found in early Hem-
ingfordian (Garvin Gully and Thomas Farm) or later local faunas of the
Gulf Coast. In addition, the presence of D. notionastes in two small
local faunas of Florida, Brooksville amd Franklin Phosphate Pit No. 2,
strengthens this view and suggests that these three faunas are similar
in age. The degree of evolution of D. notionastes at Buda and Franklin
Phosphate Pit No. 2 is advanced with respect to that found at
Brooksville, indicating that Brooksville is probably slightly older than
these two faunas.
Bassariscops achoros, as a new species, is of little value in de-
terming the age of the Buda Local Fauna, although the only other
record of this genus, B. willistoni, is from the lower part, possibly
Arikareean, of the Brown's Park Formation of Colorado (Peterson
The Nimravinae are not known to occur later than the Arikareean
elsewhere in North America (L. D. Martin, pers. comm.). The
Schizotheriinae are not known elsewhere in North America before the
late Arikareean (Skinner 1968). The overlap of these two subfamilies in
the Buda Local Fauna suggests a late Arikareean age.
The single tooth of Cynorca found at Buda has features transitional
between C. social and C. proterva. Woodburne (1969) placed this



probable transition in the late Arikareean or early Hemingfordian.
Also, Cynarctoides sp. and the phenacocoeline oreodont occur in
known western faunas no earlier than late Arikareean (Barbour and
Cook 1914; Schultz and Falkenbach 1950; Galbreath 1956; Stevens et
al. 1969). These taxa, therefore, all favor a late Arikareean age not
clearly indicated by the occurrence of Nanotragulus loomisi alone.
The Oxydactylus-like camels in the Buda Local Fauna have a tem-
poral range which extends from the Arikareean to the Barstovian in
North American faunas. These taxa, as well as the even more inclusive
group Mustelidae, provide no further refinement of the age of this


The major portion of the continental mid-Tertiary faunal record of
North America is preserved in the thin blankets of rocks that cover
much of the Great Plains from Saskatchewan to Mexico. However,
during the Tertiary Florida, as now, surely had a different spectrum of
climate, topography, soil type, and vegetation. Paleontological studies
of the Gulf Coast (Quinn 1952; Wilson 1960; Patton 1969b; Klein 1971;
Waldrop 1971; Webb 1974) indicate that the Gulf Coastal Plain has
been a distinct faunal province throughout most of the Tertiary.
The composition of the Buda Local Fauna, the first extensive
Arikareean sample in the Gulf Coastal Plain, bears a general
resemblance to Arikareean faunas of the Great Plains. Only the new
camel is completely new at the generic level. Cynorca was extremely
rare in the Great Plains during the Arikareean, as possibly was
Bassariscops, previously known only from the Brown's Park Fauna of
Most taxa in the Buda Local Fauna, while generically related to
groups found in the Great Plains during the early Miocene, are prob-
ably distinct species. Daphoenodon notionastes is distinct from the
more bear-like species of the Great Plains, but it did not survive into
Hemingfordian time when Daphoenodon superbus appears in the
Hemingfordian Garvin Gully fauna (Wilson 1960) of Texas. The Buda
Cynarctoides might be a distinct species from the Colorado and
Nebraska samples.
Nanotragulus loomisi is the only identifiable species in the Buda
Local Fauna that is not a new species. The inclusion of N. loomisi in
this local fauna constitutes a major range extension from its pre-
viously known occurrences in South Dakota and Wyoming (Lull 1922;
Loomis 1933; Frick 1937; Macdonald 1963). Nanotragulus loomisi ap-

Vol. 24, No. 2


parently retained its species identity over a range that encompassed at
least two faunal provinces, the Great Plains and the Gulf Coastal
The more indeterminate taxa in the fauna, such as the nimravine
cat, the large mustelid and the anchitherine horse, are of little interest
zoogeographically, except to record the presence of these groups in
Florida during the Arikareean.
The Buda Local Fauna is characterized by a mixture of elements
from better known faunas of western North America. Although
similarities can be seen between the Buda Local Fauna and western
faunas, the Buda Local Fauna is nonetheless distinct from Great
Plains and Pacific Coast faunas. Presumably this reflects its
geographic location in the Gulf Coastal Province where evolutionary
tendencies were influenced by wholly different conditions. Geographic
barriers helped establish species distinctions, but persistent insularity
postulated by White (1942) is not indicated in this fauna.


The Buda Local Fauna is the first extensive sample of Arikareean
mammals in Florida, and most taxa in this fauna are new to the fossil
record of Florida. Two new species are described: Daphoenodon no-
tionastes and Bassariscops achoros. A new genus of Camelidae is
recognized but not named. Other taxa are Cynarctoides sp.
Mustelidae, Nimravinae, Cynorca sp., Phenacocoelinae, Camelidae,
Anchitheriinae, Moropus sp., and Nanotragulus loomisi. Daphoenodon
notionastes, the anchitherine horse, and the two camels in the Buda
Local Fauna also occur in Franklin Phosphate Pit No. 2, another
Arikareean local fauna in Florida. The only element in common with a
third Arikareean local fauna, Brooksville, is Daphoenodon no-
Daphoenodon notionastes de-emphasized the bear-like massive
mandible and crushing dentition which are typical of related amphi-
cyonids in the Great Plains. It did not survive into the Hemingfordian
in the Gulf Coastal Plain.
Bassariscops achoros is a canid on the basis of its petrosal struc-
ture. It is recognized as a cynarctine by the presence of accessory
cusps on the carnassials. Bassariscops achoros is even more primitive
than B. willistoni in retaining wider, more angular upper molars.
Among the Cynarctinae, Bassariscops is one of the least modified
A single M, is readily referable to Cynarctoides but is the most
unspecialized specimen of that genus yet recorded.


The mustelid in the fauna is as large as Sthenictis, although generic
identification is not possible due to lack of material. The same can be
said for the identification of the nimravine felid and an anchitherine
A small species of Moropus in the Buda Local Fauna is only
broadly comparable with other small schizotherine chalicotheres from
Arikareean deposits of the John Day Basin of Oregon and Aquitanian
deposits at St. G6rand-le-Puy (Coombs 1974; in press).
An oreodont extends the range of the Phenacocoelinae into Florida
from the Arikareean of Texas (Castolon Local Fauna, Stevens et al.
Two camels in the fauna, a small form that is a new genus and a
larger, unidentifiable species, represents a group of camels with
brachydont teeth and unfused metapodials. This group had its
greatest diversity in the Gulf Coast region. The unnamed new genus is
the smallest known camel, excluding the oromerycids.
An excellent sample of Nanotragulus in the Buda Local Fauna pro-
vides sufficient examples of variation among individuals to justify
synonymizing N. "lulli" Frick and N. "intermedius" Schlaikjer with
N. loomisi, which thus had a geographic range including Nebraska,
South Dakota, and Florida.
Faunal correlations place the age of the Buda Local Fauna in the
Arikareean, probably in the late Arikareean.
The composition of the Buda Local Fauna suggests that the Gulf
Coast faunal province was as distinct from the Great Plains during the
Arikareean as later. The two new species and the unnamed new camel
genus are found only in Florida. Cynorca is found in only one, possibly
two, other localities during the Arikareean, and Bassariscops is known
only from the lower part of the Brown's Park Formation of Colorado.


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