Front Cover
 Systematic mammalian paleontol...
 Age and correlation
 Back Matter
 Back Cover

Title: Latest Pliocene mammals from Haile XV A, Alachua County, Florida
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Permanent Link: http://ufdc.ufl.edu/UF00027853/00001
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Title: Latest Pliocene mammals from Haile XV A, Alachua County, Florida
Physical Description: Book
Creator: Robertson, Jesse S.
Publisher: Florida State Museum, University of Florida,
Copyright Date: 1976
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Bibliographic ID: UF00027853
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Table of Contents
    Front Cover
        Front Cover 1
        Front Cover 2
        Page 111
        Page 112
        Page 113
    Systematic mammalian paleontology
        Page 114
        Page 115
        Page 116
        Page 117
        Page 118
        Page 119
        Page 120
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        Page 165
        Page 166
        Page 167
        Page 168
        Page 169
        Page 170
        Page 171
    Age and correlation
        Page 172
        Page 173
        Page 174
        Page 175
        Page 176
        Page 177
        Page 178
        Page 179
        Page 180
        Page 181
        Page 182
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        Page 184
        Page 185
        Page 186
    Back Matter
        Page 187
    Back Cover
        Page 188
Full Text


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of the
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Number 3






Volume 20


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SYNOPSIS: The mammalian fauna of Haile XV A is the first known from the Gulf
Coastal Plain during Blancan (Latest Pliocene) time. The mammals are repre-
sented by 8 orders, 17 families, and 18 genera, of which two are new species and
three are new to North America. Kraglievichia, a giant, extinct armadillo previously
known only from Late Miocene and Pliocene deposits of South America, is reported
for the first time in North America and a new species is described. A brief review of
the subfamily Chlamytheriinae suggests that Plaina Castellanos is a synonym of Krag-
lievichia Castellanos and that Hoffstetteria Castellanos is a synonym of Pampatherium
Ameghino. Chlamytheres are shown to have been present in North America con-
tinuously since the establishment of the late Cenozoic land bridge between North
and South America. Dasypus bellus (Simpson), a smaller extinct armadillo, is
reported for the first time in the Blancan, marking its earliest occurrence anywhere.
Glossotherium chapadmalensis (Kraglievich), a small mylodont ground sloth pre-
viously known only from the early Pleistocene of Argentina, is reported from North
America for the first time. This species may have been ancestral to both G. harlani
(Owen) and G. robustus (Owen).
The Old World Flying Squirrel, Cryptopterus, is reported for the first time in the
New World and a new species is described. The nearest related species occurs in
the Late Pliocene deposits of West Germany.
Close alliance of several Haile XV A taxa with South American Plio-Pleistocence
forms strengthens the previously suggested correlation between the Chapadmalalan
stage of South America and at least the early part of the Blancan stage of North
The abundance of aquatic non-mammalian vertebrates, together with the large
extinct otter, Satherium, and the beaver, Castor, indicate that the environment of
deposition was a permanent stream. The terrestrial community includes the tropical
or subtropical edentates Kraglievichia and Dasypus. Probable forest indicators in-
clude Mylohyus, Castor, Cryptopterus, Glossotherium, and Tapirus. The zoogeo-
graphy of several exotic forms is discussed.

' The author is an Associate Professor in the Department of Biology, Jacksonville University,
Jacksonville, Florida 32211. Most of this study was done in partial fulfillment of the Ph.D.
degree at the University of Florida. Manuscript accepted 15 July 1974.

ROBERTSON, JESSE S., JR. 1976. Latest Pliocene Mammals from Haile XV A,
Alachua County, Florida. Bull. Florida State Mus., Biol. Sci. Vol. 20(3):pp. 111-186.

5-70. 8

SG o, 3, c

SYNOPSIS --..... ------------------- ------------------------------------ 111
INTRODUCTION -....--.-----...------------------ ----- -------- 112
ACKNOWLEDGMENTS ---------.----------------- 113
GEOLOGY .........-- ------------ --------.----.- --------.----------- --- 113
SYSTEMATIC MAMMALIAN PALEONTOLOGY ---------------------------------------- 114
Order Insectivora
Cryptotis parva ..----.--------------------------------- 114
Scalopus aquaticus _--_------.------..--------...-------- 115
Order Edentata
Glossotherium chapadmalensis --..-........--- --.---- ---------- 116
Kraglievichia new species .........------------------.- ------ 124
Review of the Chlamytheriinae -...--------------- ---------------- 135
Dasypus bellus .......------...........-------------------------- ------- 142
Order Lagomorpha
Sylvilagus sp. ----....------------------------ 146
Order Rodentia
Cryptopterus new species -..--.-...-.......-------- 147
Castor canadensis ---------...----------- ---------- 153
Sigmodon medius ....---.------------ ----------------- -- 153
. Order Carnivora
Canidae .....----. .... ... ... -----------.---------------------- 154
Satherium sp. ...---------- ---------------- ------- 154
Smilodon gracilis ..-----..--- --------- ----.-------- 157
Order Proboscidea
Gomphotheriidae .-----------.---- ------ ------------ 158
Order Perissodactyla
Nannippus phlegon --....---.... -------------- ---- 158
Equus (Dolichohippus) simplicidens ....-..---------. ----------.------ 158
Equus (Hemionus) calobatus -..-..- -- - ------- ------- ---- 160
Tapirus sp. ---..--------------..------ --------- 163
Order Artiodactyla
Mylohyus floridanus ----.----..-----........ ------- ---- 163
Hemiauchenia cf. macrocephala .----------------------------- ------ 167
Odocoileus virginianus .-......---------------------.---.-------------.. 169
AGE AND CORRELATION --_.--..-.........-.--------..----------------------------- 170
PALEOECOLOGY ..----..... ----------------------------- ---------- ----------- 176
ZOOGEOGRAPHY --.........-----.---.---.---------.--------.------ ------------178
SUMMARY --------..... .....----------.---------------. 179
BIBLIOGRAPHY ----....------------- -------- ---.-------- -- 181


Florida has long been noted for its rich Late Pleistocene fossil verte-
brate deposits, including the Vero site in Indian River County (Weigel
1962), the Melbourne Bone Bed in Brevard County (Gazin 1950, Ray
1958), Seminole Field in Pinellas County (Simpson 1930), Reddick (Ray
et al. 1963) in Marion County, and many others. In recent years a few


sites have yielded fossil vertebrates of older Pleistocene and even Latest
Pliocene age. These Blancan sites are of particular interest, inasmuch as
they constitute the first samples from eastern North American (Webb
1974a). This report is the first systematic study of one of the Latest
Pliocene sites.
The Haile XV A locality is situated on the property of Parker Brothers
Limestone Products Incorporated, near Haile, T9S, R17E, Section 25,
S NW 1/4 of SW 1/4, Alachua County, Florida. In 1964 the site was dis-
covered and some specimens collected by Phillip Kinsey of Jacksonville
SBeach, Florida. Further excavations were carried out in the same year
S by S. David Webb, Robert Allen, and myself with the support of National
Science Foundation Grant GB 3862.

Fossil material was examined from the American Museum of Natural History
(AMNH), the University of California, Museum of Paleontology (UCMP), the
Florida State Museum (UF), the University of Houston (HCT), the University of
Kansas (UK), and the United States National Museum of Natural History (USNM).
I wish to thank the persons in charge of these collections for the opportunity to
study the materials in their care.
I also wish to thank S. David Webb, Thomas H. Patton, and H. K. Brooks for
their encouragement and help during the course of this study.

The Haile XV A deposit lies at an elevation of about 90 ft above
sea level and is a filled fissure in the Ocala (Eocene) Limestone. This
fissure measures approximately 10 m along its east-west axis, and its
north-south dimensions range irregularly from about 3 to 5 m (Fig. 1).
The bottom of the fissure-filling sequence lies at a depth ranging from
1 to 7 m below the land surface. It consists of a stratum of dark brown,
compact, iron-stained clays (or "hardpan"), with a thickness ranging from
2 to 6 m.
The fossil-bearing matrix overlies the "hardpan," and consists of an
alternating sequence of sands and clays. The lowermost unit is a coarse
gravelly sand, containing calcareous cement and concretions. It is the
thickest unit in the fossiliferous sequence, ranging from 2 to more than
6 m in depth, and is the most productive fossiliferous horizon. Many of
the included larger bones are encrusted with cemented sand and gravel.
The clay units are pure, greenish in color, massive in texture, and vary
from 10 to 20 cm. The sand layers decrease in thickness upward through
the sequence, with the uppermost sands being less than 1 m thick. The
S clays vary little in thickness, except that the uppermost unit in the fos-
siliferous section is nearly a meter thick.


I C.l0 i mtors D
0 1 1 1

FIGxrE 1. Geologic section at Haile XV A: (A) sandstone alternating with gray
clay; (B) massive greenish clay; (C) coarse, poorly sorted, gravelly sandstone; (D)
dark brown hardpan; and (E) Ocala Limestone (Eocene).
In several instances fossil bones in a sand layer have been found pro-
jecting partly into the overlying clay layer. In each instance the pro-
jecting part of the bone shows no noticeable wear or weathering. Evi-
dently each clay unit was deposited rapidly over the preceding sand unit,
with little or no intervening period of erosion. Overlying the fossiliferous
sequence of sands and clays is a layer of thin, brown sandstone that is
nonfossiliferous and grades upward into the soil profile.

Twenty mammalian taxa, representing a minimum of 34 individuals,
were collected at Haile XV A (Table 1).
Cryptotis parva (SAY 1823)
MATERIAL.-UF 17466, right mandible with M1-M3.
This specimen is indistinguishable from Recent Florida specimens of
C. parva, except for its slightly larger size (Table 2). The only other
Blancan record of Cryptotis is from the Rexroad fauna (Hibbard 1937,
1941), Meade County, Kansas.

Vol. 20, No. 3



Species MNI
Cryptotis parva 1
Scalopus aquaticus 2
Glossotherium chapadmalensis 1
Kraglievichia floridanus 3
Dasypus bellus 1
Sylvilagus sp. 3
Cryptopterus webbi 1
Castor canadensis 1
Sigmodon medius 2
Canidae 2
Satherium sp. 1
Smilodon gracilis 1
Gomphotheriidae 1
Nannippus phlegon 2
Equus (Dolichohippus) simplicidens
Equus (Hemionus) cf. calobatus 2
Tapirus sp. 1
Mylohyus floridanus 1
Hemiauchenia cf. macrocephala 1
Odocoileus virginianus 5

UF 17466 AND OTHER FOSSIL AND RECENT Cryptotis parva.

C. parva1 (20) C. parva' (fossil)
UF 17466 X OR X2 OR3
Condyloid to mental foramen 6.8 6.1 5.9-6.7 6.6 6.1-7.0 (8)
Condyloid to M1 6.7 6.9 6.7-7.4 7.1 6.8-7.6 (8)
Condyloid to M2 5.3 5.7 5.5-6.0 5.8 5.5-6.1 (8)
MI-M2 2.8 2.6 2.4-2.7 2.7 2.5-2.9 (8)
M1-M3 4.0 3.4 3.3-3.6 3.6 3.5-3.6 (2)
M2-M3 2.5 2.2 2.1-2.4 2.3 2.2-2.3 (2)

SMeasurements after Martin (1974).
3 0. R.=observed range; number of specimens in parenthesis

Scalopus aquaticus LINNAEUS 1758

MATERIAL.-UF 17467, 2 left M2; UF 17465, right humerus.
A comparison of the three specimens from Haile XV A with corre-
sponding elements of modern S. aquaticus from the southeastern United
States reveals no significant differences in size or morphology.

Scalopus is also present in the Blancan Rexroad fauna of southwestern
Kansas (Hibbard 1941) and in the Sand Draw local fauna of north-
central Nebraska (Hibbard 1972). Another mole (Hesperoscalops)
from the Rexroad fauna is based on a lower dentition. A direct com-


prison could not be made with that genus, because only upper teeth are
known from Haile XV A. As Hibbard (1941) points out, however, mole
humeri are quite diagnostic at the generic level, so it seems unlikely that
a Scalopus-like humerus could belong to any other genus.

Glossotherium (Glossotherium) chapadmalensis (KRAGLIEVICH 1925)
Eumylodon chapadmalensis Kraglievich 1925
Glossotherium (Glossotherium) chapadmalensis (Kragl.) Hoffstetter 1952
MATERIAL.-UF 10922, partial skeleton.
The Haile XV A specimen probably represents one individual, be-
cause there is no duplication of elements and the left and right elements
are very similar in all measurements. Most, if not all specimens were
taken from the basal sand layer of the fossiliferous sequence, and some
parts of the skeleton and dermal ossicles were nearly articulated.
CRANIUM AND UPPER DENTITION.-The upper dentition of the Haile
XV A specimen (Fig. 2) is complete except for the left upper caniniform.
The right upper caniniform is triangular in cross-section and very well
developed. The stoutness of the caninform is reflected by great trans-
verse expansion of the anterior portion of the maxilla, as is also true of
the holotype from Argentina. This contrasts strikingly with G. harlani,
which has a reduced or absent caniniform and transversely narrow
The first upper molariform tooth is oval in cross-section. As in G.
harlani and G. robustus, it is the longest anteroposteriorly of all the upper
teeth but is relatively narrow in transverse diameter. Because the well-
developed lower caniniform occludes in part with this tooth, wear on the
anterior oblique surface in G. chapadmalensis is much more accentuated
than in G. harlani. In G. robustus specimens observed, this tooth was
worn off smoothly just above the alveolus. A very short diastema sepa-
rates the upper caniniform tooth from the first molariform tooth in the
specimen from Haile XV A.
The second upper molariform tooth of UF 10922 is triangular in cross-
section. The lingual side is the narrowest and contains a deep inflection.
The transverse diameter is greater than the anteroposterior diameter in
both the Florida and the Argentina specimens of this species, whereas
the opposite is true in G. harlani. Of the two G. robustus specimens for
which measurements are provided, one resembles G. chapadmalensis,
whereas the two diameters in the other specimen are nearly equal. The
third molariform tooth of UF 10922 is similar in shape to the second and
morphologically similar to the corresponding teeth in both G. harlani

Vol. 20, No. 3


FIGURE 2. Glossotherium chapadmalensis: upper dentition X 0.85.

and G. robustus. The fourth molariform is relatively small compared to
the Argentina specimen of G. chapadmalensis, although it is identical in
The palate is very constricted in the area of the last upper teeth and
considerably expanded at the anterior end of the maxilla. Both of these


characters are more extreme in UF 10922 than in the holotype or in speci-
mens of G. harlani and G. robustus.
The remainder of the cranial material consists of part of the occipital
bone and portions of the left and right periotic bone. The occipital frag-
ment consists of the left condyle and a portion of the otic region. The
hypoglossal canal, jugular foramen, and auditory meatus show no observ-
able differences from those of G. harlani and G. robustus. A portion of
the left squamosal bone is attached to the occipital. Part of the right
squamosal is present, including the zygomatic process.
MANDIBLE AND LOWER DENTITION.-The lower caniniform of UF 10922
(Fig. 3, 4) is well developed and directed dorsolabially. This tooth
is relatively larger than the corresponding tooth in the South American
specimen of G. chapadmalensis. The tooth has a chisel-like tip, the result
of dual occlusion with the upper caniniform and first molariform teeth.
The second molariform tooth is rectangular and has its long axis
oblique to the tooth row. Stock (1925) measured the long axis as the
anteroposterior length; Kraglievich (1925) used the shorter axis of the
tooth. For purposes of comparison, Kraglievich's method is used here
for the Haile XV A specimen and Stock's method for measuring G.
robustus. (For consistency one must reverse the measurements of G.
robustus provided by Kraglievich [1928].) It is obvious from Table 3
that the orientation of the teeth, not their basic morphology, probably
accounts for the interspecific differences in this tooth.
The third lower molariform tooth is of a slightly different shape in
G. chapadmalensis. The two main columns are separated by a thin
bridge, while in G. robustus this bridge is thicker.
Unfortunately, the mandibular symphysis is not preserved in the Haile
XV A specimen, but the portion of the mandible present is similar to
that of the other two species.
VERTEBRAE.-The atlas of UF 10922 is the only well preserved verte-
bra. According to Stock (1925) the atlas of G. harlani differs from that
of G. robustus in the more posterior position of the lateral process. The
location of the dorsal foramina also varies; in G. harlani they are well
separated as compared to those in G. robustus. In the expression of
these two characters, UF 10922 resembles C. harlani.
LIMB BONEs.-The right humerus and both ulnae are present, as are
the left femur, both tibiae, and the left fibula. Hind foot bones repre-
sented are the astragalus, calcaneum, and various metatarsals and pha-
langes. The small size of this species is indicated by the measurements
of these limb bones (Table 4).
Several bones of the manus warrant a brief description. The cunei-
form appears to be somewhat different in G. chapadmalensis than in G.

Vol. 20, No. 3


FIGURE 3. Glossotherium chapadmalensis: right mandible, lateral view, X 0.60.


FIGURE 4. Glossotherium chapadmalensis: right mandible, occlusal view, X 0.60.

Vol. 20, No. 3


G. chapadmalensis G. harlani G. robustus
Haile XV A Argentina1 California2 Argentina3
Anteroposterior length, tooth 1 14.7 14.0 17.9 17.9
Transverse length, tooth 1 10.6 10.5 16.0 18.2
Anteroposterior length, tooth 2 22.4 22.0 33.3 22.3
Transverse length, tooth 2 12.0 15.0 17.4 16.7
Anteroposterior length, tooth 3 15.6 20.5 27.1 22.8
Transverse length, tooth 3 22.4 24.0 23.0 23.0
Anteroposterior length, tooth 4 17.9 15.0 22.6 20.3
Transverse length, tooth 4 20.5 24.0 23.8 22.4
Anteroposterior length, tooth 5 21.4 24.0 24.3 23.3
Transverse length, tooth 5 13.6 18.0 19.4 19.2

Anteroposterior length, tooth 1 20.5 14.5 21.4 20.4
Transverse length, tooth 1 12.4 10.5 16.0 15.0
Anteroposterior length, tooth 2 18.7 18.0 28.3 23.8
Transverse length, tooth 2 14.5 22.0 22.5 22.8
Anteroposterior length, tooth 3 14.5 14.0 28.8 25.0
Transverse length, tooth 3 20.5 25.0 18.2 19.0
Anteroposterior length, tooth 4 40.3 43.0 51.9 50.2
Transverse length, tooth 4 17.3 18.0 23.8 20.2

'Measurements after Kraglievich (1925).
SMeasurements after Stock (1925).
RMeasurements after Owen (1842).


TABLE 4.-LIMB MEASUREMENTS (IN MM) OF Glossotherium chapadmalensis AND
Glosotherium harlani.

G. chapadmalensis G. harlanil
UF 10922, Haile XV A Rancho La Brea
left right
Greatest anteroposterior distance
of head 70.0 70.0 123.8
Greatest width of shaft at deltoid
ridge 71.3 122.6
Width of distal articular surface 86.0 133.2
Greatest length 285 395.9
Width of distal articular surface 34 55.0
Total length 355 546.4
Transverse diameter of head 79.3 127.8
Least width of shaft 99.5 164.6
Greatest width across distal
tuberosities 140 234.8
Width of distal condyles 111 188.5
Width of intercondyloid space 27.1 47
Width of inner condyle 47.2 47.2 88.7
Vertical extent of inner condyle 67 69.8 120
Total length 181 176 247.3
Greatest width, proximal end 114 185.2
Greatest width, distal end 90 88 142.5
Anteroposterior distance, distal
end 64.8 66.6 101.9
Total length 183 263
Width, proximal end 50.5 103.6
Width, distal end 39.5 73.5

'After Stock (1925). The measurements represent the mean of large samples. The number of
specimens varied from element to element and in some cases was not given.

harlani; in the UF specimen it has a square palmar outline, whereas in
Stock's figure (1925: Fig. 72d) of G. harlani it is rectangular. This dif-
ference is reflected by the measurements of the cuneiform in Table 5.
In UF 10922 the proximal-distal dimension is greater than the distance
across the ulnar articular surface, while the opposite is true for 39 speci-
mens measured by Stock. The cuneiform appears to be a deeper element
in G. chapadmalensis than in G. harlani. The remaining elements of the
manus preserved in the Haile XV A specimen are the left scaphoid, right
pisiform, left metacarpal III, and left phalanx II, digit III.
DIscussIoN.-This specimen so closely resembles Eumylodon ( = Glos-
sotherium) chapadmalensis that it is here ascribed to that species. G.

Vol. 20, No. 3


Glossotherium chapadmalensis FROM HALE XV A AND Glossotherium

G. chapadmalensis
left right G. harlani'

Greatest distance across articular
surface 46.2 75.3(39)2
Greatest distance across dorsal
surface from inner side to outer 42.1 68.5(42)
Greatest proximal-distal distance 34.3 50.5
Distance across articular surface
for ulna 31.2 61.3
Greatest length 25.3 55.6(22)
Greatest depth 18.9 36.4
Greatest width 20.9 34.7
Greatest length 71.7 103.1(30)
Width, proximal end 51.9 75.2
Width, distal end 30.5 50.9
Depth, proximal end 42.0 66.0
Length 40.5 48.9(42)
Depth, inner condyle 23.5 39.9
Width, proximal end 26.2 47.9
Greatest width, anterior end 57.4 91.8(21)
Greatest depth 83.4 125.4
Anteroposterior diameter 76.1 140.2(41)
Length 84.3 119.3(29)
Width, proximal end 41.2 41.5 60.7
Depth, proximal end 34.3 37.1 45.5
Width, distal end 38.5 50.1
Depth, distal end 35.3 43.5
Length 30.2 31.0 35.4(32)
Depth, inner condyle 20.4 21.5 21.8
Width, proximal end 22.6 22.5 35.1
Length 104.4 174.1
Proximal-distal distance of ungual
base 41.0 73.5
Width, proximal end 32.4 55.0
1 Measurements after Stock (1925).
2Number in parentheses refers to the sample size from which the mean (the numbers in the
right-hand column) was calculated.


chapadmalensis was described from the Chapadmalal fauna (Early
Pleistocene) of Argentina and is here reported from North America for
the first time. It is a small species that appears to be very closely re-
lated to both G. harlani and G. robustus, but it differs from these two
species in the nature of the anterior dentition and overall size. It re-
sembles both in the morphology of most of the postcranial elements, with
minor exceptions in the manus and pes. G. chapadmalensis is thus tem-
porally, geographically, and morphologically a plausible ancestor of both
G. harlani and G. robustus.
Much confusion has arisen concerning proper usage of the names
Mylodon and Glossotherium. In his thorough review of the literature,
Kraglievich (1928) concluded that Mylodon darwinii is the genotype of
Mylodon, and the species described by Owen (1840) as Mylodon ro-
bustus should actually be included in a distinct genus, Glossotherium.
However, Kraglievich retained Paramylodon (Brown 1903) as a valid
genus, even though Stock (1925) included it in the synonomy of Mylodon
harlani. Simpson (1945) correctly argued that if the North and South
American Late Pleistocene forms are not generically distinct, they should
all be referred to the genus Glossotherium. Hoffstetter (1952) retained
Paramylodon as a subgenus of Glossotherium, this distinction being
based on the narrow muzzle, the long cranium, and frequent absence of
the upper caniniform tooth. Following this usage, the Haile XV A
species should be called Glossotherium (Glossotherium) chapadmalensis.

Kraglievichia floridanus new species

HOLOTYPE.-UF 10902, partial skull collected by P. E. Kinsey, S. D. Webb,
R. R. Allen, and J. S. Robertson in 1964.
TYPE LOCALITY AND HORuzON.-Haile XV A, T9S, R17E, Sec. 25, NW 1/4 of
SW 1/4, Alachua County, Florida, Blancan.
DIAGNOSIs.-Differs from Kraglievichia paranensis in having the fourth upper
tooth reniform rather than peg-like, with the long axis anterolingual instead of parallel
to the tooth row.
REFERRED MATERIAL.-UF 10902, mandible and postcranial elements (probably
the same individual as the type skull); UF 17474, right ulna, also from Haile XV A;
UF 10432, right humerus, Santa Fe I; UF 9354, right humerus, Santa Fe I; UF
10830, right radius, Santa Fe I; UF 16371, left metatarsal IV, Waccasassa River;
UF 17475, right metatarsal IV, Santa Fe II; UF 17472, right metatarsal III, Santa
Fe II; UF 17476, right femur, Haile XIII; UF 17568 and UF 17569, right naviculars,
Haile XV A.

DESCRIPTION.-The Haile XV A skeleton and isolated postcranial
material from other sites in Florida provide the first adequate samples
of the postcranial skeleton of this genus.
The following description is based primarily on the Haile XV A speci-

Vol. 20, No. 3


men; but because certain elements are lacking, it has been supplemented
by material from other Florida Blancan localities.
CRANUM.-The nasal bones (Fig. 5b) of UF 10902 compare favor-
ably in size and morphology with those of the South American specimen
of Kraglievichia paranensis described by Castellanos (1927). They
differ, however, in having tiny protuberances on the anterior ends near
the medial surface. These structures are not present on any of the other
specimens or figures of Pampatherium or Kraglievichia studied. Only
the anterior halves of the nasals are present in UF 10902, because the
skull had been eroded away, and only parts of it recovered.
The premaxilla (Fig. 5a) contains only one alveolus. The premaxil-
lary-maxillary suture forms the posterior border of the first alveolus (or
tooth socket), as in Pampatherium. Holmesina (from North America)
was originally thought to be distinguished by the presence of a single
premaxillary tooth; two such teeth were supposed to occur in Pampa-
therium. A single alveolus occurs in the neotype of "Holmesina"
(AMNH 26856; Simpson 1930) as well as in UF 889 (P. septentrionalis).
However, it now appears that the true Pampatherium also has only one
tooth in the premaxilla (G. Edmund, pers. comm.), and, therefore, the
supposed difference between the North and South American forms is
probably not valid.
The major features of the maxilla from Haile XV A compare favorably
with those in Pampatherium. The infraorbital foramen (Fig. 5e) is
located directly above the sixth tooth in both genera. The anterior
palatal foramina (Fig. 5c) are located between the posterior edges of
the fourth teeth in the Haile XV A specimen, whereas their position
varies somewhat in Pampatherium. The maxillary process of the zygo-
matic arch lies directly above the seventh tooth in both Kraglievichia
and Pampatherium.
Only a portion of the zygomatic arch is present in UF 10902 (Fig.
5d-e). The zygomatic process of the maxilla turns posteriorly and down-
ward as it leaves the skull. The anterior portion of the jugal, which
borders the zygomatic process laterally, then turns upward and expands
posteriorly to accept the squamosal process. A well-developed suture,
located at the posterior end of the jugal, indicates that the zygomatic
arch is complete in Kraglievichia, as it is in Pampatherium. The base
of the zygomatic process of the maxilla is expanded by sinuses, as in
UPPER DENTITION.-There are nine teeth in both the upper and
lower jaws of Kraglievichia. One of the upper teeth is located in
the premaxilla. (Because the incisors of most mammals occur in the
premaxilla, it is tempting to refer to the first tooth in Kraglievichia as an




7 a

26 nvn

FIGURE 5. Kraglievichia floridanus, cranial material: (A) right premaxillary; (B)
nasal; (C) palatal portion of maxillary; (D) zygoma, ventral view; (E) zygoma,
lateral view; (F) right mandible, occlusal view; (G) right mandible, lateral view;
(H) first lower tooth; (I) fourth lower tooth. Abbreviations: iof=infraorbital
foramen, ju = jugal, mx = maxillary, mx-pmx = premaxilla, n = nasals, numbers refer to
tooth sequence.

Vol. 20, No. 3


incisor; however, I have continued to use the non-committal terminology
"tooth one" through "tooth nine" in this report.) The fourth tooth pro-
vides critical evidence for the phylogenetic position of the Haile XV A
species. The alveolus for this tooth suggests that it was incipiently bilo-
bate and nearly parallel with the more posterior teeth, a condition inter-
mediate between earlier Kraglievichia and later Pampatherium.
Measurements of the cranium and upper dentition are presented in
Table 6.
MANDIBLE AND LOWER DENTITION.-Partial left and right mandibles
are preserved in UF 10902. In the right mandible (Fig. 5f-g) the 7th
tooth is complete, the 8th is broken, and alveoli for the 6th and 9th teeth
are present. Unfortunately, the anterior part of each mandible is lacking
in UF 10902. Other than size, no differences are apparent between Krag-
lievichia and Pampatherium with regard to the posterior parts of the
mandibles (see Table 6).
Vertebrae.-The vertebral material from the Haile XV A skeleton
consists of 7 thoracic, 5 lumbar, and 4 caudal vertebrae. Five of the
thoracic vertebrae are cemented together in the proper sequence by
coarse sandstone. Comparison of these specimens with the vertebrae
of a well-preserved skeleton of Pampatherium from Branford IA, Suwan-
nee County, Florida (Rancholabrean), shows no significant morphologi-
cal differences except for size.
FRONT LIMBs.-In this description the major elements of the forelimb
are compared with those of both Dasypus and Pampatherium. Kraglie-
vichia floridanus is approximately the same size as Rancholabrean speci-
mens of Dasypus bellus (a large extinct Pleistocene armadillo) and con-
ceivably the limb elements of these two species could be confused. The
podial elements and hind-limb elements should never be confused, how-
The right humerus of UF 10902 is well preserved, except for the distal
end (Fig. 6a-b). The humerus of Kraglievichia is greatly expanded
laterally (as seen in the specimens from Santa Fe I). The supracondy-
lar foramen is relatively larger than in Dasypus. The articular surface
for the radius is concave in Kraglievichia, whereas it has a slight con-
vexity in Dasypus. The supinator ridge is relatively narrow in Kraglie-
vichia, and the deltoid ridge is wider. Comparison of the proximal end
of the humerus in Kraglievichia and Pampatherium shows no distinct
differences, except for size.
Three ulnae are known from the Haile XV A site; two of which are
probably from the same individual because they are similarly preserved,
are from opposing sides, and agree closely in measurements (Table 7).
The ulna of Kraglievichia (Fig. 6c-d) is laterally flattened and has a long


Kraglievichia Kraglievichia Pampatherium Pampatherium
UF 10902 AMNH 26856 UF 889, 890
Haile XV A Argentina1 Florida Florida
Width of nasals at anterior border of premaxilla 25.9 27.6 39.0
Width of palate at center of tooth 4 15.0 17.4 28.02 29.6
Anteroposterior length, tooth 4 12.3 12.6 16.7
Anteroposterior length, tooth 5 15.4 12.7 23.7 20.5
Anteroposterior length, tooth 7 16.4 15.0 22.6 23.6
Width, tooth 1 6.8 6.5 7.6
Distance, top of ninth alveolus to bottom of jaw 37.8 32.8 60.2 51.4
Length, tooth 1 5.4
Width, tooth 1 3.7
Length, tooth 3 8.2 12.3
Width, tooth 3 5.2 6.9
Length, tooth 6 (alveolar) 16.0 14.8 25.1 25.1
Length, tooth 7 14.3 13.9 23.3 22.1
Width, tooth 7 7.1 11.2 9.2
Width, tooth 8 (alveolar, anterior lobe) 11.9 10.5 10.0
Length, tooth 9 10.3 13.8
Width, tooth 9 5.5 7.8

SMeasurements after Castellanos (1927).



FIGURE 6. Kraglievichia floridanus, front limb elements: (A) right humerus, medial
view; (B) right humerus, anterior view; (C) left ulna, medial view; (D) left ulna,
anterior view; (E) left radius, posterior view; (F) left radius, anterior views; X 0.45.


UF 10902, HAILE XV A.

Left Right

Lateral width, proximal end 36.7
Anteroposterior width, proximal end 36.8
Greatest anteroposterior diameter of shaft 25.5
Lateral width of shaft at same location 18.2
Distance from proximal end to top of entepicondylar foramen 100.5
Total length 129.0
Lateral width, proximal end 17.2
Lateral width, distal end 16.1
Lateral width at semilunar notch 25.5 25.7
Anteroposterior width at semilunar notch 29.4 30.5
Total length 89.2 87.2
Lateral width, proximal end 22.6 22.3
Anteroposterior width, proximal end 11.7 11.4
Lateral width, distal end 23.0 22.3
Anteroposterior width, distal end 17.4 16.5
Total length 32.8
Width, proximal end 10.9
Depth, proximal end 14.5
Width, distal end 10.5
Depth, distal end 11.2
Total length 34.4 36.3
Width, proximal end 13.8 14.5
Depth, proximal end 12.8 13.0
Width, distal end 12.6 13.4
Depth, distal end 10.5 11.0
Total length 29.1
Width, proximal end 11.4
Depth, proximal end 10.3
Total length 193.5
Lateral width, proximal end 60.9
Anteroposterior thickness of greater trochanter 41.7
Lateral width of shaft at third trochanter 41.3
Anteroposterior thickness of shaft at same point 23.7
Greatest lateral width of articular facets 48.4
Greatest anteroposterior width, distal end 51.5
Total length 121.0
Lateral width, distal end 46.7
Anteroposterior width, distal end 25.1

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Total length 63.3
Width of articular facets for astragalus 28.0
Depth of facet for cuboid 12.2
Width of facet for cuboid 11.5
Greatest lateral width 39.5
Greatest anteroposterior distance 23.3
Greatest depth 22.5
Total length 33.2 32.8
Width, proximal end 12.2 12.5
Depth, proximal end 13.8 14.3
Width, distal end articularr surface) 13.2 13.7
Depth, distal end articularr surface) 11.3 10.2
Total length 36.5 34.5
Width, proximal end 15.0 14.9
Depth, proximal end 14.7 14.4
Width, distal end articularr surface) 13.2 12.3
Depth, distal end articularr surface) 11.3 11.5
Total length 24.6
Width, proximal end 17.6
Depth, proximal end 10.7

olecranon process. The articular facets for the radius and medial condyle
of the humerus are combined in this form, whereas in Dasypus they are
partially divided. Another striking difference is a lateral groove that runs
the entire length of the ulna in Dasypus and terminates at the upper
border of the semilunar notch in Kraglievichia. Size appears to be the
only difference between the ulna of Kraglievichia and Pampatherium.
In both Kraglievichia and Dasypus the distal end of the radius is
massive compared to the proximal end, although it is less flattened in
Kraglievichia (Fig. 6e-f). The proximal portion of the shaft is relatively
thicker and less curved in Kraglievichia. Comparison with Pampather-
ium shows only a size difference.
The left metacarpal II is present in the Haile XV A skeleton, and
generally agrees with that of Pampatherium in shape and proportion.
The facet for articulation with the trapezoid, however, shows an im-
portant difference; when viewed laterally, this facet is smoothly curved
in Pampatherium and has a sharp V-shape in Kraglievichia.
The magnum facet is oblong in Pampatherium and round in Krag-
lievichia. The trapezoid facet is wider dorsally in Pampatherium. To


compensate for this, the facet for metacarpal III bulges out proximally
to form a pocket for the trapezoid facet. In Kraglievichia less of the
trapezoid articulates with metacarpal III.
The left and right third metacarpals are preserved in UF 10902 from
Haile XV A and also in UF 10722 from Santa Fe I. In this element the
facet for metacarpal II and the trapezoid shows a relatively greater as-
sociation with the trapezoid than with the adjacent metacarpal. In both
Pampatherium and Kraglievichia the magnum facet is convex dorsally
and concave ventrally. In Kraglievichia the greater portion of the facet
is convex, whereas the opposite is true in Pampatherium. When viewed
from below, the ventral portion of this facet lies oblique to the long axis
of the bone, whereas in Pampatherium it is perpendicular to the long
HIND LIMBS.-The femur (Fig. 7a-b), the tibia, and the fibula (Fig.
7c-d) show no morphological differences from those of Pampatherium
except for size.
The calcaneum of Kraglievichia is less expanded distally and less
robust than that of Pampatherium (Fig. 8). In Kraglievichia the two
astragalar facets are connected to form a bilobed facet, whereas in Pam-
patherium they are separated by a central valley. In Kraglievichia the
facets are nearly equal in size, whereas in Pampatherium the lateral
facet is much the larger of the two.
There are no apparent differences in the naviculars of Kraglievichia
and Pampatherium. It may be noted that the three right naviculars
from Haile XV A exhibited little variation in the relative shapes and
sizes of the facets in which the cuneiform bones articulate.
In metatarsal II the mesocuneiform facet is relatively more narrow
ventrally in Kraglievichia than in Pampatherium. There is an indenta-
tion at the proximal end, at the point where it meets the proximal end
of the metatarsal I; in Pampatherium there is no such indentation, the
medial border forming an unbroken line. The metatarsal I of Kraglie-
vichia presumably had a corresponding projection; no such feature oc-
curs in metatarsal I of Pampatherium.
In lateral view the proximal articular surface of metatarsal III appears
rounded in Kraglievichia, while in Pampatherium it forms a straight line
perpendicular to the long axis of the element. Because the proximal end
of this element is rounded in Kraglievichia, the articular surface can also
be seen when the element is viewed from above. This would appear to
permit more dorsoventral movement of the toes of Kraglievichia.
The facets for metatarsal III and the cuboid are united on metatarsal
IV in Kraglievichia, but separate in Pampatherium.
Two ungual phalanges of digit V (UF 10902) have been recovered

Vol. 20, No. 3


FIGURE 7. Kraglievichia floridanus, hind limb elements: (A) right femur, anterior
view; (B) right femur, lateral view; (C) right tibia and fibula, anterior view; and
(D) right tibia and fibula, lateral view; X 0.45.

FIGURE 8. Kraglievichia and Pampatherium, calcanea, anterior views: (A) Kraglievichia floridanus, right calca-
neum, Haile XV A; (B) Kraglievichia sp., right calcaneum, Inglis IA; and (C) Pampatherium septentrionalis,
right calcaneum, Branford IA; X 0.80.


from Haile XV A; one (presumably from the forefoot) is rather narrow
and pointed, and the other (presumably from the hind foot) blunt and
broad. They agree in all features except size with the corresponding
phalanges in Pampatherium. Measurements of the limb element in
other examples of Kraglievichia from various Florida localities are pre-
sented in Table 8.

The genus Kraglievichia is reported here for the first time in North
America. In order to place the genus in a meaningful context, the sub-
family Chlamytheriinae is reviewed briefly. The nomenclatural history
of the various chlamythere genera will be discussed, followed by a brief
characterization of the various genera, beginning with the oldest.
The first remains of a chlamythere were discovered in a Brazilian
cave deposit in 1836 by Peter Wilhelm Lund and described by him as
Chlamytherium humboldtii (Lund 1838). Lund consistently used his
original generic name in his early works, but later (beginning about
1840) he emended this to Chlamydotherium, calling the earlier name a
lapsus. Several later authors followed this secondary spelling. Bronn
(1838) meanwhile had given the name Chlamydotherium to a genus of
glyptodonts. Ameghino (1875) proposed Pampatherium as a substitute
for Lund's supposedly preoccupied generic name, Chlamydotherium,
but he later discontinued use of the new name after checking the
spelling in Lund's original description. Paula Couto (1956) has revised
the use of Pampatherium, which has come back into general usage.
The first North American record of Pampatherium was reported by
Leidy (1889a), although he first designated the new species Glyptodon
septentrionale. Leidy (1889b) later referred the same material to the
South American species Chlamytherium humboldtii. Sellards (1915)
believed that the North and South American forms represented different
species and resurrected Leidy's original specific name septentrionale.
Ameghino (1902) described Machlydotherium from the Eocene of
Patagonia. This still stands as the earliest record for a chlamythere.
Castellanos (1927) named two new genera of chlamytheres: Vassallia,
based on an edentulous mandible and several dermal plates; and Krag-
lievichia, based on two skulls, a mandible, and a small amount of post-
cranial material. The type species of Vassallia is Chlamytherium minu-
tum (Moreno and Mercerat 1891). Kraglievichia was erected to include
C. paranesis, C. intermedia (Ameghino 1887), and C. subintermedius
Rovereto 1914), with C. paranensis as the generic type.
Simpson (1930) established the genus Holmesina, including in it the



UF 10432 (right) UF 9354 (right)
Humerus Santa Fe I Santa Fe I
Greatest anteroposterior diameter of shaft 28.8
Lateral width of shaft at same point 19.5
Lateral width, distal end 60.2 54.2
Greatest width of distal articular surface 37.8 36.8

UF 10449 (left) UF 17474 (right)
Ulna Santa Fe I Haile XV A
Total length 116.5
Lateral width, proximal end 17.6
Lateral width, distal end 10.4
Lateral width at semilunar notch 20.5 25.1
Anteroposterior width at semilunar notch 23.4 31.8
UF 10830 (right)
Radius Santa Fe I
Total length 105.2
Transverse width, proximal end 24.5
Anteroposterior width, proximal end 13.3
Transverse width, distal end 23.1
Anteroposterior width, distal end 20.6

UF 17476 (left)
Femur Haile XII B
Lateral width, proximal end 55.3
Anteroposterior width, proximal end 34.2
Lateral width of shaft at third trochanter 38.8
Anteroposterior width of shaft at same point 16.5
UF 17473 (left)
Calcaneum Santa Fe II
Total length 64.1
Width of articular facets for astragalus 24.5
Depth of facet for cuboid 14.9
Width of facet for cuboid 10.6

UF 17568 (right) UF 17569 (right)
Navicular Haile XV A Haile XV A
Greatest lateral width 37.3 33.6
Greatest anteroposterior distance 22.6 18.3
Greatest depth 30.6 26.6
UF 10902 (left)
Metatarsal II Haile XV
Total length 30.8
Width, proximal end 11.3
Depth, proximal end 13.6
Width, distal end articularr surface) 12.3
Depth, distal end articularr surface) 10.2

Vol. 20, No. 3



UF 17472 (right)
Metatarsal III Santa Fe II
Total length 34.2
Width, proximal end 16.5
Depth, proximal end 15.2
Width, distal end articularr surface) 13.5
Depth, distal end articularr surface) 11.4
UF 17475 (right)
Metatarsal IV Santa Fe II
Total length 29.5
Width, proximal end 11.2
Depth, proximal end 12.2
Width, distal end articularr surface) 15.1
Depth, distal end articularr surface) 10.5

species septentrionalis, after studying well-preserved material from the
Seminole Field in western peninsular Florida. Subsequent authors
tended to refer the North American forms to Holmesina and those from
South America to Chlamytherium.
Castellanos (1937) later proposed a new genus, Plaina, the type
species of which (C. intermedius) he had earlier placed in Kraglievichia.
His rationale for establishing this new genus was based largely on his
interpretation of the lineage of the chlamytheres. Castellanos believed
that logically there should be a form intermediate in size between Krag-
lievichia and Chlamytherium, and because C. intermedius is larger than
the other material referred to Kraglievichia, he saw it as representing this
intermediate form.
The genus Hoffstetteria Castellanos (1957) was based upon a skull
collected in Ecuador, which had been described previously as a new
species, C. occidentalis, by Hoffstetter (1952).
Machlydotherium is the oldest of the various chlamythere genera, its
remains being known from the Eocene of Patagonia. Its precise re-
lationship to later chlamytheres is not clear (Simpson 1945), a situation
that may be attributed partly to the absence of any Oligocene records of
the subfamily.
Vassallia occurs in Araucanean (Pliocene) deposits in Argentina
(Castellanos 1946) and is also known (a skull; UCMP 40401) from the
Late Miocene La Venta fauna of Colombia. This identification is based
upon the nature of the anterior dentition (the first five teeth are peg-like
and rotated lingually) and the small size of the specimen.
Porta (1962) reported Kraglievichia from the La Venta fauna of
Colombia, but this record was based only upon dermal plates. It now
seems more likely that they represent Vassallia instead.


Castellanos (1927) listed two major morphological differences be-
tween Vassallia and Kraglievichia. In Vassallia the first five teeth tend
to be peg-like and rotated lingually, whereas in Kraglievichia only the
first four teeth show this condition. Vassallia is also significantly smaller
than Kraglievichia. I suspect here, as did Castellanos, that Vassallia is
the ancestor of Kraglievichia.
So far as the status of Plaina is concerned, Castellanos (1927) was
probably correct in his initial judgment when he assigned the material
on which this genus was based (three isolated dermal plates) to the
genus Kraglievichia. He considered these plates to be intermediate in
size and sculpturing between Kraglievichia and Pampatherium, al-
though they actually fall within the smaller size range of Kraglievichia.
Furthermore, it seems illogical to interpret slight variations in plate rugo-
sity as generically significant. I therefore propose that Plaina be re-
garded as a synonym of Kraglievichia.
The genus Hoffstetteria Castellanos (1957) is based upon supposed
differences in shape and measurements of the teeth. These differences
are minor, however, and may be attributed to specific variation. Hoff-
stetteria is thus considered to be a synonym of Pampatherium.
In Simpson's (1930) description of Holmesina, he listed a number of
characters by which this new genus differed from Pampatherium.
James (1957) subsequently showed that the characters given by Simpson
are not sufficient to separate the North and South American forms generi-
cally, and that Holmesina is a synonym of Pampatherium. I support
James' conclusion. Simpson (1930) argued also that if Holmesina was
not valid, then all South America forms should be placed in the genus
Pampatherium, as they are no more different from each other than is
Pampatherium from Holmesina. This argument is not without merit.
However, new characters discovered during the present study, consid-
ered together with those previously recognized, provide ample justifica-
tion for recognition of Kraglievichia and Pampatherium as distinct
To summarize, the genera of chlamytheres recognized here are:
Machlydotherium (Eocene, South America); Vassallia (Miocene and
Pliocene, South America); Kraglievichia, including Plaina (Pliocene,
South America and Plio-Pleistocene, North America); and Pampather-
ium, including Hoffstetteria and Holmesina (Pleistocene, North and
South America).
The most obvious morphological trend in chlamythere evolution since
the Miocene has been a consistent increase in size. Vassallia is slightly
smaller than Kraglievichia, which in turn is substantially smaller than

Vol. 20, No. 3


Pampatherium. This increase in size may be seen by comparison of the
two femora in Figure 9.
Inasmuch as the increase in size noted above occurred gradually,
many intermediate stages logically would be expected. A series demon-
strating the changes in size between Kraglievichia and Pampatherium
appears in Figure 10. The dermal plates illustrated are from various
Plio-Pleistocene stages in Florida. Plates from the Early Blancan Haile
XV A site are the smallest, followed by those from the Early Irvingtonian
Inglis IA site. Still larger plates occur in the later Irvingtonian Coleman
IIA site, and the largest plates of all are those from the Rancholabrean
Branford IA locality.
Another major trend in chlamytheriine evolution is the shift from
peg-like to bilobate tooth shape and from an oblique to a parallel orienta-
tion. This change seems to have progressed in an anterior direction,
inasmuch as an increase in the number of anteriorly-situated bilobate
and parallel teeth may be seen in the transition from Vassallia through
Pampatherium (Castellanos 1937).
The Haile XV A specimen differs from previously described speci-
mens of Kraglievichia and Pampatherium in the nature of the anterior
upper dentition. In South American specimens of Kraglievichia, the first
four teeth are oval in cross-section rather than reniform and have their
long axes turned lingually. In Pampatherium the first three teeth show
this condition, but the fourth is usually bilobate and parallel with the
rest of the tooth row. In James' (1957) specimen the first four teeth
are oval, but the long axis of the fourth tooth parallels the rest of the
tooth row, rather than being canted lingually. The peg-like appearance
of the fourth tooth is presumably a primitive condition that recurred
in an aberrant individual. The incipiently bilobed condition in the
fourth upper tooth of Kraglievichia floridanus represents the transitional
condition between Kraglievichia and Pampatherium, although the fea-
tures of the feet remain primitive in several respects.
Chlamytheres apparently originated in South America, because all
pre-latest Pliocene records of this group are restricted to that continent.
The oldest genus is the Eocene Machlydotherium. No chlamytheres are
known from the Oligocene, but Vassallia is present in Miocene deposits,
and both Vassallia and Kraglievichia occur in the Pliocene of South
America. Kraglievichia is here reported in Blancan deposits of North
America. Pampatherium is present throughout the Pleistocene in both
North and South America. This distribution presents some interesting
geographic and phylogenetic problems.
Until now the evolution of the chlamytheres was thought to have
occurred exclusively in South America, with only the end product,



FIGURE 9. Pampatherium and Kraglievichia, femora, anterior views: (A) Pam-
patherium septentrionalis, right femur, Branford IA; and (B) Kraglievichia flor-
idanus, right femur, Haile XV A; X 0.45.

Vol. 20, No. 3


Pampatherium, having migrated to North America during Ranchola-
brean time. Study of the Haile XV A fauna and other pre-Ranchola-
brean sites of Florida indicates instead that Kraglievichia migrated to

FIGURE 10. Kraglievichia and Pampatherium, dermal plates, dorsal views: (A)
Kraglievichia floridanus, Haile XV A; (B) Kraglievichia sp. Inglis IA; (C) Pam-
patherium septentrionalis, Coleman IIA; and (D) Pampatherium septentrionalis,
Branford IA; X 0.30.


North America shortly after the establishment of the Late Cenozoic land
bridge between the two continents, and that chlamytheres evidently
continued to evolve in Florida and the Gulf Coastal Province throughout
the Pleistocene.
Three hypotheses can be offered regarding the Pleistocene zoogeo-
graphy and evolution of the chlamytheres. The first suggests that from
a common Kraglievichia origin North and South American Pleistocene
chlamytheres had separate but parallel histories. If so, Holmesina would
be the correct name for the later North American chlamythere.
A second hypothesis suggests that Kraglievichia spread from South
America into North America near the end of the Pliocene, after which it
became extinct in South America. This would account for the absence
of chlamythere records in the Early Pleistocene of South America. (A
dermal plate that was questionably attributed to the Chapadmalalan
stage by Kraglievich [1934] actually came from a locality near the major
deposit, and its horizon is unknown. Since each chlamythere shell pro-
duces a high number of bony plates, they are usually abundant if present
at all.) Later in the Pleistocene, then, Pampatherium spread back into
South America.
The third hypothesis suggests that chlamytheres were continuously
distributed through tropical America during most of the Pleistocene.
Thus the evolution of Pampatherium from Kraglievichia was a single
continuous event that spread through populations from the Gulf Coastal
region and mesoamerican parts of North America into the northern parts
of South America. The absence of Early Pleistocene deposits in tropical
South America makes it difficult to test this hypothesis. This last hy-
pothesis, here favored, may be partly tested by careful comparisons
of later Pleistocene chlamytheres.

Dasypus bellus SIMPsON 1929
MATERIAL.-UF 16698: left nasal, right maxilla and mandible, right astragalus,
and 51 isolated dermal plates from Haile XV A; UF 10449: right ulna from Santa Fe
The dentigerous specimens probably represent one individual, be-
cause they articulate closely. Most of the plates also were found closely
associated. The present material has been compared with other speci-
mens of Dasypus bellus and with its closest living relative, Dasypus
novemcinctus. The only other known D. bellus specimens containing
teeth are the excellent skeleton from Crankshaft Pit, Missouri (UK 15544;
Oesch 1967), and a fragmentary jaw from Melbourne (Ray 1958). Com-
parisons have been made with postcranial material from numerous Pleis-
tocene sites in Florida.

Vol. 20, No. 3



FIGURE 11. Dasypus bellus, mandible, UF 16698, Haile XV A: (A) occlusal view;
and (B) lateral view; X 1.45.

The nasal bone is represented by the anterior portion only and is
larger than, but morphologically similar to the nasal bone of D. no-
In discussing the dentition, teeth will be designated as follows: T1
will refer to the first upper tooth, T2 the second upper tooth, etc. Simi-
larly numbered subscripts will be used in referring to lower teeth.
The maxilla is broken posteriorly and contains only T3-T6. Except
for size, the lateral portion of this specimen compares closely with the
corresponding position of the Crankshaft Pit specimen. In the Haile
XV A specimen T3 is transversely flattened, but all the rest are round
and peg-like. The preceding two teeth of the Haile XV A Dasypus (UF
16698) were probably also transversely flattened, as this is the usual con-
dition in D. bellus. In D. novemcinctus the first three teeth tend to be less
flattened laterally than in D. bellus. The remaining upper teeth were
probably round and peg-like as is usually the case with T4-T6. Talmage
and Buchanan (1954), as well as others, have pointed out that tooth
number and structure are variable in Dasypus. In four D. novemcinctus
skulls examined, the total number of teeth varied from seven to nine.
In UF 16698 (Fig. 11), as in the Kansas mandible, two foramina are
present-one large and one small. In UF 16698 the larger of the two
foramina occurs between T2 and Ts. The smaller of the two foramina
is located 38 mm anterior to T1. In UK 15544, the larger foramen is
anterior to the smaller and is located directly beneath T_. The smaller
is located beneath Ts. The number of these mental foramina in D.
novemcinctus varies from one to four. The symphysis of the Haile XV A
specimen is weak, as is characteristic of the genus.
The mandible of UF 16698 lacks T4-Ts. The preceding teeth, T,-Ts,
are laterally flattened, but the succeeding teeth, T, and T,, are round in


UF 16698 UK 15544
Haile XV A Crankshaft Pit
Florida Missouri
left right

Total distance, teeth 3-6 19.9 23.81 23.7
Anteroposterior distance, tooth 3.5 4.61 4.5
Transverse distance, tooth 3 2.1 4.61 4.5
Anteroposterior distance, tooth 4 4.4
Transverse distance, tooth 4 3.6
Anteroposterior distance, tooth 5 4.0 4.1
Transverse distance, tooth 5 4.2 5.1
Anteroposterior distance, tooth 6 4.2 5.22 4.5
Transverse distance, tooth 6 4.1 4.8
Depth of ramus at tooth 1 6.8 12.4
Width of ramus at tooth 1 2.8 3.7
Depth of ramus at tooth 6 8.4 15.2 15.5
Width of ramus at tooth 6 4.8 8.2 7.8
Depth of ramus at tooth 8 7.2 13.5 12.2
Width of ramus at tooth 8 5.2 7.1 6.4
Length of tooth row 38.8 45.6
Anteroposterior distance, tooth 1 3.0 2.83
Transverse distance, tooth 1 1.3 1.83
Anteroposterior distance, tooth 2 3.3 2.2 2.1
Transverse distance, tooth 2 1.4 2.7 2.3
Anteroposterior distance, tooth 3 3.3 4.0 4.5
Transverse distance, tooth 3 2.2 3.2 3.2
Anteroposterior distance, tooth 4 4.81 4.7 4.6
Transverse distance, tooth 4 2.81 4.5 4.2
Anteroposterior distance, tooth 5 4.01 4.1 4.4
Transverse distance, tooth 5 3.32 4.5 4.5
Anteroposterior distance, tooth 6 4.5 4.3 4.5
Transverse distance, tooth 6 3.7 4.7 4.7
Anteroposterior distance, tooth 7 4.6 4.6 4.2
Transverse distance, tooth 7 3.5 4.7 4.6
Anteroposterior distance, tooth 8 4.1 3.5 3.4
Transverse distance, tooth 8 4.2 3.5 3.4
1 Alveolar.
2Alveolar, estimated.
SNot completely erupted.

cross-section. In this feature the Haile XV A specimen differs from the
lower dentition of UK 15544, in which all the teeth are round. Except
for smaller size and this tendency toward flattening of the anterior teeth,
the mandibles and dentition of the Haile XV A specimen differ little from
other Dasypus material. Dental measurements appear in Table 9.
In the ulna from Santa Fe I the channel on the medial side is deep
and persists to the proximal end of the element. This distinguishes it
from the otherwise very similar ulna of Kraglievichia (p. 127).

Vol. 20, No. 3


The astragalus of the Haile XV A D. bellus, along with other speci-
mens of D. bellus from Florida Pleistocene sites, differs little from Recent
specimens of D. novemcinctus, except for size. A comparison of the
astragali of Dasypus from various stages of the Pleistocene reveals a pro-
gressive size increase similar to that of the chlamytheres.
The dermal plates of UF 16698 do not differ from any other speci-
mens of Dasypus, except for size. Martin (1974:41) measured samples
of plates from several Florida sites and showed little overlap in plate size
between D. bellus and D. novemcinctus. His measurements indicate
that the size of the plates does not follow the trend through the Pleisto-
cene that is evident in other parts of the D. bellus skeleton. The Cole-
man IIA plates are notably small and overlap slightly in size with those
of D. novemcinctus. However, this is probably a function of the small
sample size used; there are approximately 2500 plates of diverse sizes
in a single armadillo carapace, and there is no satisfactory method for
determining from which parts of the shell the sample may have come.
Size trends based on plates thus should involve large samples in order
to insure accuracy.
The morphological differences between the Haile XV A material and
later D. bellus material are very slight and presumably have a variational
basis. Consequently, the Haile sample is referred to Dasypus bellus.
Dasypus bellus has been considered a southern North American
species of Rancholabrean age. However, Martin (1974) extended the
temporal range of the species back into Irvingtonian time, and the pres-
ent study places the earliest record in the Blancan.
A question arises regarding the relationship between D. bellus and
D. novemcinctus. These animals are clearly more closely related to each
other than to any other armadillos (Auffenberg 1957), and it is uncertain
whether D. novemcinctus was derived from D. bellus or existed allo-
patrically with that species during the Pleistocene, replacing it during
the last few thousand years. If D. novemcinctus did evolve from D.
bellus, it appears to have done so quite rapidly. In Miller's Cave (Pat-
ton 1963) a date of ca 8000 BP was given for the Travertine stratum
bearing D. bellus remains, as compared to ca 3000 BP for the overlying
Brown Clay deposits containing D. novemcinctus. Thus, the change oc-
curred in that area within a span of only 5000 years. Another possibility
is that D. novemcinctus existed somewhere else during the Pleistocene
and replaced D. bellus very late in the Pleistocene. If so, it would be
expected earlier in some Central or South American Pleistocene deposits.
To date, the only fossil record of D. novemcinctus is from Miller's Cave,
and the only Late Pleistocene armadillos from Mexico are referable to


D. bellus. Until more tropical fossil records are found, the problem
must remain unresolved.
In view of the long residence of Dasypus bellus in Florida, it seems
surprising that no pre-Rancholabrean records are available elsewhere in
North America. Possibly Dasypus was restricted to the Gulf Coastal
region during most of the Pleistocene. Slaughter (1961) cited two
ecological conditions that may have been necessary for this species: (1)
winters no more severe than those occurring in North-central Texas
today, and (2) rainfall more than 20 inches per year. Thus the distri-
bution of such climatic conditions may have restricted D. bellus to Flor-
ida and the Gulf Coastal Plain during most of the Pleistocene.

Sylvilagus sp. GRAY 1867

MATERIAL.-UF 17561: 2 M3; UF 17562: M3; UF 17563: lumbar vertebra;
UF 17564: 3 innominates; UF 17565: 2 femora; UF 17566: 1 tibia; UF 17567:
2 metatarsals.

Hypolagus, Alurolagus, and Sylvilagus are the smaller lagomorphs
common in North American Blancan deposits. Unfortunately, M3 and
Ms, the only teeth preserved in the Haile XV A fauna, are not diagnostic
(Dawson 1958, Downey 1970). When compared with the Hypolagus
innominates (Dawson 1958), the Haile XV A specimens possess a less
elevated ilial crest, which is probably a more advanced condition. The
remainder of the Haile XV A material resembles Sylvilagus in size and
morphology, but one cannot completely discount reference of this ma-
terial to Alurolagus (Downey 1968, 1970) from the Benson and Curtis
Ranch Faunas of Arizona.
A species determination is not possible from the available material.
Two living species of Sylvilagus occur widely in Florida, S. palustris and
S. floridanus. The genus is presently distributed throughout North
America, and also ranges into South America where it is represented by
S. floridanus and S. brasiliensis (Hall and Kelson 1959).
Previously, the principal Blancan records of Sylvilagus were from the
Broadwater fauna (Barbour and Schultz 1937) and the Sand Draw fauna
(Skinner and Hibbard 1972) of Nebraska. Numerous Irvingtonian and
Rancholabrean records of this genus are also known throughout North

Vol. 20, No. 3


Cryptopterus webbi new species1

TYPE.-UF 12353: partial right mandible with third molar.
TYPE LOCALrrY.-Haile XV A, T9S, R17E, Section 25, NW 1/4 of SW 1/4,
Alachua County, Florida.
DIAGNOSIS.-Large sciuropterine; low-crowned, subovate M. not tapered posteri-
orly; protoconid connected to weak anteroconid; large anterolabial cingulum as in
Cryptopterus tobieni; flat trigonid basin bearing "chaos" of low oblique ridges; strong
metaconid and metaconid-metastylid crest; distinct entoconid; faint hypolophid; broad
posterolophid bearing vestigial hypoconulid; posterolabial flexid absent.

DESCRIPTION.-The unique flying-squirrel specimen from Haile XV A
consists of the posterior part of the right mandible bearing a moderately-
worn third molar. Much of the ascending ramus, especially the condyle,
is well preserved. The distance between the last molar and the condyle
is about 15 mm. A large dental foramen lies 6.1 mm posterior to the last
molar. The angular region is massive, concave lingually, and descends
to a depth of more than 12 mm below the alveolar level, at which point
it is broken. The coronoid region is also broken.
The third molar measures 4.91 mm in length, 3.70 mm maximum
width across the metaconid and protoconid, and 3.29 mm posterior width
across the entoconid and hypoconid. This is by far the largest flying
squirrel in the New World, rivaling the extinct Eurasian species of
Cryptopterus and Petauria and some living Asian species of Petaurista.
The Haile XV A molar has a subovate outline and is only slightly
tapered toward the posterior end. This readily distinguishes it from
specimens of Miopetaurista and Pliopetaurista, in which the posterior
end of the tooth tends to be long and narrow. The crown is low, the
protoconid rises only 1.1 mm above the crown base; it thus contrasts with
most species of Petaurista and especially with Eupetaurus (McKenna
On the labial side of the molar the strong protoconid and hypoconid
are separated by a moderately well-developed mesoconid. The meso-
conid is more robust than in Pliopetaurista and does not produce a labial
spur, such as Sulimski (1964) noted in the specimens from W6ze, Poland,
The protoconid is joined to a small anteroconid by a short, anteriorly
curved crest, and thence weakly to the prominent metaconid. The weak
anteroconid does not extend toward the labial wall of the molar as in

1 Named for Professor S. David Webb in honor of his contributions to the vertebrate paleontol-
ogy of Florida.


FIGURE 12. Cryptopterus webbi type, right mandible, anterior end upwards, UF
12343, Haile XV A; X 5.40.

Vol. 20, No. 3


4 mm

FIGURE 13. Cryptopterus webbi, right M,, anterior end upwards, UF 12353, Haile
most sciuropterines. Instead, a large cingulum occupies the anterolabial
corner of the molar, recalling the arrangement found in the third lower
molar of Cryptopterus tobieni (Mein 1970: fig. 39).
The anterolingual part of the third molar is greatly expanded as in
most sciuropterines. It supports a very large prominent metaconid from
which a posterior crest connects broadly back to a low metastylid (or
mesostylid according to Mein's 1970 terminology), and a labial crest
reaches the anteroconid. From the robust metaconid, the enamel surface
slopes gradually downward into a very shallow trigonid valley. In its
center are several low sinuous ridges (lophulids) that trend in a postero-
lingual direction. Such lophulid complexes are characteristic of many
sciuropterines; the homologous but higher ridges in Petaurista xanthotis
were appropriately termed "metaconid-metastylid chaos" by McKenna
(1962). These lophulids presumably provide drainage canals for the
juices produced when pulpy foods are crushed in the trigonid basin.
Similar features for similar purposes are found in such unrelated mam-
mals as Ailuropoda and Pongo.
A distinct entostylid occupies the midlingual part of the molar from
Haile XV A. A weak ridge, the hypolophid, extends labially from the
entostylid, but is soon lost in the broad continuity between the trigonid
and talonid "valleys" in the center of the tooth. A minor depression,


separating the entoconid from the strong posterolophid, encircles the
posterior margin of the tooth. A thickening of the enamel in the middle
of the posterolophid evidently represents a vestigial hypoconulid. Al-
though faint, this structure is notable inasmuch as Mein (1970:22)
characterized all other Cryptopterus as having a smooth posterolophid
without any hypoconulid. There is no sign of a posterolabial inflection
in the molar from Haile XV A.
DIscussioN.-Although the affinities of the Haile XV A flying squirrel
might reasonably be sought among other North American sciuropterines,
such comparisons prove unsatisfactory. The North American record
consists of Glaucomys of Pleistocene and Recent age and a late Miocene
sample of two small species from the Cuyama Valley of California de-
scribed as Sciuropterus by James (1963), but assigned to Cryptopterus
by Mein (1970). James (1963) suggested that the North American Mio-
cene species might have been ancestral to Glaucomys, or at least that no
closer ancestry is known. Mein (1970) was more skeptical of a direct
relationship. In either event the Haile XV A specimen, far from bridging
that gap, differs in a number of basic features from what would be ex-
pected of a Latest Pliocene Glaucomys ancestor. Whereas Recent
Glaucomys is even smaller than the Cuyama Miocene fossils, the Haile
XV A specimen is much larger. Neither an anteroconid nor an antero-
lingual cingulum (of any size) occurs in either the California Miocene
specimens or in Glaucomys, yet both features are found in the Haile
XV A specimen. The protolophid directly connects the protoconid to
the metaconid in both the Miocene Cuyama specimens and in Recent
Glaucomys specimens, whereas it is interrupted by an anteroconid in
the Haile XV A specimen. The talonid valley is not crenulated in the
Haile XV A specimen, and the trigonid lophulids are heavier than in
other North American flying squirrels.
Turning to the Old World fauna, we find a much greater variety of
both fossil and Recent genera with which to make comparisons. These
include several taxa whose size equals that of the Haile XV A specimen,
and some that exceed it. It seems evident that the phylogenetic de-
velopment of the flying squirrels was centered in Eurasia, and it is fortu-
nate that a relatively complete fossil record of the group is known from
Europe. Mein (1970), who recently published the most complete
phylogenetic review of the sciuropterine squirrels, recognized three broad
groups of fossil and Recent genera based on lophule development and
enamel crenulation. His arrangement is as follows:
GROUP I (Enamel smooth; lophules absent)
Recent Genera: Glaucomys, Eoglaucomys, lomys.
Fossil Genera: Cryptopterus, Petauria.

Vol. 20, No. 3


GROUP II (Enamel smooth; lophules present)
Recent Genera: Pteromys, Trogopterus, Ptero-
myscus, Belomys, Aeretes, Petaurista, Eupetaurus.
Fossil Genera: Miopetaurista, Forsythia, Pliope-

GROUP III (Enamel crenulated; lophules rare or absent)
Recent Genera: Petinomys, Hylopetes, Aeromys.
Fossil Genera: Blackia, Pliopetes.

Of these diverse genera, European fossil specimens of Cryptopterus
and Petauria bear the closest resemblance to the specimen from Haile
XV A. As in those genera, the enamel in the Haile specimen is not
finely crenulated. The pattern of cuspids and lophids agrees closely
with that in some species of Cryptopterus. Unfortunately, the question
of whether the lophules in the upper cheek teeth were elaborately de-
veloped cannot be directly answered by the Florida specimen, but if
the detailed resemblances of Cryptopterus lower molars are correct, loph-
ules would be absent from the upper molars just as they are from the
Several features of the lower molar from Haile XV A rule out its
relationship to Group II genera. The nearly ovate shape of the last
molar is a reliable distinction from molars of the extinct taxa Miope-
taurista and Pliopetaurista. These taxa also bear a distinct posterolabial
flexid that is barely (if at all) recognizable in the Florida molar. As
Sulimski (1964) noted in his material from Poland, the mesoconid ex-
tends labially as a spur in Pliopetaurista, in contrast to the round meso-
style in the Florida specimen. Finally, the metastylid (mesostylid of
Mein 1970) is very prominent in Miopetaurista and Pliopetaurista,
whereas in the Florida specimen it is largely submerged by the broad
crest connecting it to the metaconid. Each of these features indicates a
closer relationship of the Haile XV A specimen with Group I taxa.
The flying squirrel tooth from Haile XV A most closely resembles the
third lower molars of Cryptopterus. It also resembles homologous teeth
of Petauria helleri from Early Pleistocene red earth fissure fillings in the
Solenhofen Limestone near Schernfeld, Bavaria (Dehm 1962), but that
genus completely lacks a mesoconid on its lower molars, and the hypo-
lophid is more strongly developed than in the Florida specimen. This
general resemblance of the Florida specimen to the Bavarian specimen
led to its preliminary assignment to "Petauria sp." in Webb (1974a). It is
now evident, however, that the Haile XV A specimen is perhaps more
accurately recognized as a new species of the genus Cryptopterus.


When one compares Cryptopterus webbi with other species of that
genus, the closest resemblance is to those from the very Late Pliocene.
In view of the Blancan age of the Florida species, this is what one might
expect. The youngest European species of Crytopterus is C. tobieni
from the lignites of Wolfersheim-Wetterau, West Germany (late Perpi-
gnan equivalent). The slightly older species, C. thaleri, resembles C.
tobieni in size and many other features but is not presently represented
by any lower molars. Special features shared by lower third molars of
C. tobieni and C. webbi are the broad anterolabial cingulum, the weak,
anteriorly-directed anteroconid, the short protolophid, the mestastylid
little distinct from the metaconid ridge, the distinct entoconid, and the
weakly developed hypolophid. C. webbi is slightly smaller than C. tobieni
and perhaps C. thaleri, and it differs from all other known Cryptopterus
specimens in the presence of a vestigial hypoconulid. Lower third molars
of the Miocene species of Cryptopterus differ from those of C. webbi and
C. tobieni in having a narrow anterolabial cingulum, a strong labially
directed anteroconid, a complete protolophid, a more distinct metastylid,
and the entoconid attached both to the posterolophid and to a strong
hypolophid. An exception to this is the presence of a distinct entoconid
and a weak hypolophid in Gaillard's (1899) original figured specimen
from La Grive, which is now referred to C. gaillardi (Mein, 1970).
Evidently this late Miocene variant embodied some of the progressive
features that were developed more definitely during the Pliocene in C.
tobieni and C. webbi. Cryptopterus mathewsi from Late Miocene de-
posits in the Cuyama Valley of California also differs in some of these
same features from C. webbi and C. tobieni; however, the lower third
molar is poorly known (James 1963). No lower dentition has been re-
covered in C. uphami material from the same area. The anteroconid is
essentially absent, as is the anterior cingulum. I am not yet convinced,
in fact, that the Cuyama species are referable to Cryptopterus.
The itinerary by which Cryptopterus reached Florida in the Blancan
must remain in doubt, but the general pattern of distribution may be
deduced. Since the nearest affinities of C. webbi are with C. tobieni
(Late Pliocene of Europe), it seems probable that the genus reached
North America from the Old World during that epoch. We may safely
say that the date of arrival in North America was within late Hemphil-
lian or Blancan time. Furthermore, the only likely dispersal route was
via the Bering Land Bridge between Asia and North America. Flying
squirrels do not truly "fly," and their present distribution patterns do not
indicate great vagility. This New World occurrence of Cryptopterus
indicates continuity of subtropical rain forest from Eurasia into eastern
North America during the Late Pliocene.

Vol. 20, No. 3


C. canadensis C. canadensis
UF 17489 Recent
Haile XV A N X OR
Width, distal end 38.7 6 36.6 34.1-38.4
Anteroposterior depth, distal end 29.2 6 29.7 28.5-32.5
Width, external condyle 13.5 5 14.0 12.2-15.5
Width, internal condyle 12.5 5 12.1 11.6-12.5
Width of shaft at third trochanter 25.2 6 29.1 27.0-30.5
Anteroposterior diameter of shaft at third
trochanter 13.9 6 11.5 10.2-13.5

Castor canadensis LINNAEUS 1758

MATERIAL.-UF 17489: left femur.

This element, which lacks the proximal end and represents a young
individual, shows no morphological differences from the living species.
A comparison of measurements (Table 10) with examples of young,
Recent Castor canadensis shows no significant size differences. Other
Blancan records of Castor in North America are from the San Joaquin
locality in California (Kellogg 1911, Stirton 1935) and the Hagerman
fauna of Idaho (Stirton 1935, Zakrzewski 1969).

Sigmodon medius GIDLEY 1922

MATERIAL.-UF 17489: left Ml; UF 12341: right MI; UF 12337: left M2; UF
12339-12340: right M3; UF 12334, UF 12338: left M,; UF 12336: left M2; UF
12342: right MI and M3 (both unworn).

This species characteristically possesses only two or three roots on the
M,. If accessory roots are present, they are centrally located and are
very small, peg-like structures. The labial root is always better devel-
oped than the lingual. The two specimens from Haile XV A (UF 12338)
exhibit these characteristic features.
Sigmodon medius is nearly identical morphologically with a closely
allied species, Sigmodon minor. According to Martin (1974), the only
difference between the two species, other than size, is that the reentrant
folds are deeper and narrower in S. minor than in S. medius. This char-
acter is not obvious in the relevant Haile XV A specimens because they
show significant wear. Discrimination between these two species must


TABLE 11.-DENTAL MEASUREMENTS (IN MM) OF Sigmodon medius AND Sigmodon

S. medius S. minor
Length M1 2 2.25 2.18-2.32 45 1.89 1.72-2.19
Width Mi 2 1.24 1.23-1.25 48 1.31 1.17-1.48
Length M2 1 1.49 -49 1.40 1.22-1.62
Width M2 1 1.53 46 1.36 1.20-1.50

SRe-calculated from Martin (1970).

depend on size alone. The larger measurements of the Haile XV A
material (Table 11) indicate that it belongs to S. medius.
Sigmodon medius is one of the most common small rodents in the
Blancan of North America; its occurrences include the Benson local fauna
of Arizona (Gidley 1922), the Vallecito fauna of California (Downs and
White 1968), the Hudspeth and Red Light local faunas of Texas (Strain
1966, Akerston 1970), and the Sand Draw fauna of Nebraska (Skinner
and Hibbard 1972).


MATERIAL.-UF 17492-17493: 2 tibiae.

Two poorly preserved tibiae indicate the probable presence of a small
canid. They are about the size of a fox, but positive identification is
not possible as only the much worn proximal ends are preserved.

Satherium sp.

MATERIAL.-UF 17487: humerus; UF 17491: right metatarsal II; UF 17490:
right metatarsal III; UF 17494: medial phalanx.

The humerus and podial elements closely resemble those of both
Pteronura, the living giant, flat-tailed otter of South America, and Sather-
ium, the extinct giant otter, from various Blancan sites in North America.
Despite the numerous close resemblances between these two genera and
the probable ancestral relationship of Satherium to Pteronura, I follow
Gazin (1934) and Bjork (1970) in recognizing the extinct genus as dis-
tinct. These two giant genera are surely more closely related to one
another than to Lutra.
The humerus from Haile XV A (Fig. 14) is more robust than are

Vol. 20, No. 3


FIGURE 14. Satherium sp., humerus, anterior view, UF 17487, Haile XV A; X 1.95.


Satherium Pteronura
Hagerman, Id. Haile XV A brasiliensis Lutra canadensis
USNM 23266 UF 17487 Recent, South America Recent, North America
Greatest width, distal end 33.4 33.0 3 36.3 33.5-40.1 7 25.1 24.1-27.2
Greatest width, distal condyle
(measured from anterior side) 21.0 21.0 3 23.3 21.5-26.5 7 15.1 11.8-17.3 >
Length of entepicondylar foramen 5.5 5.8 3 5.9 5.4- 6.5 6 4.4 3.7- 4.7
Width of entepicondylar foramen 3.3 3.9 3 4.2 4.0- 4.4 6 2.0 1.5- 2.3
Greatest anteroposterior thickness
of shaft 13.5 17.4 3 16.1 14.7-18.5 7 13.1 11.3-14.8
Transverse width at same location 9.9 10.3 3 10.2 9.5-11.4 7 7.9 7.5- 8.8
Greatest anteroposterior thickness
of distal condyle 15.4 16.1 3 14.6 12.7-16.2 7 11.2 10.2-12.4


Haile XV A Port Kennedy
UF 17496
Length 41.6 48
Width 45.2 38
Width of trochlea 28.7 27
Vertical diameter of head 18.7 17
Transverse diameter of head 26.6 25
External elevation of trochlea 21.1 22
SMeasurements after Cope (1899).

humeri of Satherium from the Hagerman local fauna of Idaho (Bjork
1970). In this respect the Haile specimen agrees more closely with speci-
mens of living South American Pteronura (Table 12). In the dimen-
sions of the metatarsals and in most other respects, however, the three
samples are remarkably homogeneous. The distal width of the Haile
humerus likely would be greater were it not for the pathological condi-
tion of the medial side; the bone has turned distad and narrowed and
is associated with much osteoporotic tissue. This anomaly does not seem
to have affected either the articular surface nor the entepicondylar fora-
men, which are located laterally and proximally, respectively, to the
affected area. The most impressive features that distinguish the Haile
XV A specimen from the Hagerman skeleton (USNM 23266) are the
much greater medial dimensions of the shaft and the deeper and wider
ectepicondylar ridge. In all respects, however, there are strong similari-
ties among the Haile sample and the genera Satherium and Pteronura.
Smilodon gracilis COPE 1880
MATERIAL.-UF 17496: right tibia (distal end) and right astragalus; UF 17498:
right metacarpal II.
The Haile material presumably is all from a single medium-sized
animal. All three bony elements were found together, the tibia and
astragalus articulate well, and the material is similarly preserved. The
material is referred to the genus Smilodon on the basis of two characters
(see Merriam and Stock 1932): (1) the presence of the astragalar fora-
men; and (2) the medial facet for the calcaneum and the facet for the
navicular are merged.
The Haile XV A sabercat was very small and was similar in size to
Smilodon gracilis. A comparison of the measurements of the astragalus
from Haile XV A with those of the astragalus of S. gracilis from the Port
Kennedy Bone Bed (Cope 1899) shows a close correlation in size
(Table 13). Inasmuch as S. gracilis is the only small species of Smilodon


known from North America and is of relatively early Pleistocene age, the
Haile XV A specimen is referred to that species.
Two species of small sabercats are known from South America.
Smilodon cruciens is known only from a single mandible (Ameghino
1904). Smilodontidion riggsi (Kraglievich 1948), which is slightly larger
than the Haile XV A form, is present in the Chapadmalal fauna and is
known only from postcranial material.1 The astragalus of this specimen
is preserved but in poor condition, and the critical characters of the facets
cannot be verified from the figures. Although other species occur both
in the Chapadmalal fauna and the Haile XV A fauna, it does not follow
that the two cats are necessarily related. Nonetheless, the overall faunal
resemblance, together with the similarity in morphology of the South
American samples, suggest that possible synonymy of the Chapadmalalan
form with Smilodon gracilis ought to be considered.

MATERIAL.-UF 17464: a small portion of a molar and two small pieces of ivory.

The complicated nature of the cheek tooth clearly indicates that this
specimen is a gomphotheriid. Further identification is not possible be-
cause of the inadequate material available.

Nannippus phlegon (HAY 1899)
MATERIAL.-UF 17484-17485: both upper cheek teeth; UF 17547: metapodial;
UF 17548: proximal phalanx; UF 17549: four ungual phalanges.

A nearly unworn upper cheek tooth 51 mm high indicates that this
form was strongly hypsodont. The other well-worn upper tooth (Fig.
15) and the postcranial material agree in all respects with the typical
Blancan species, N. phlegon. Measurements are provided in Table 14.

Equus (Dolichohippus) simplicidens COPE 1892
MATERIAL.-UF 10909: partial skull; UF 17556: partial maxilla with deciduous
molariform tooth; UF 10894: 15 cervical vertebrae; UF 10877: 37 thoracic verte-
brae; UF 10895: 13 lumbar vertebrae; UF 10896: 2 sacral vertebrae; UF 10898:
3 humeri (2 left, 1 right); UF 10919: 1 right radius; UF 10910: 2 left pelves; UF
10911: 4 femora (2 left, 2 right); UF 10915: 2 right metatarsals; UF 10921: 9
splints; UF 10916: 4 cuboids (2 left, 2 right); UF 10917: 3 calcanea; UF 10920:

' Churcher's (1967) cautionary note suggesting that the specimen may have leaked down from
younger deposits is noted.

Vol. 20, No. 3


2 astragali; UF 10917: 4 proximal phalanges; UF 10913: 4 distal sesamoids; UF
10914: 7 ungual proximals.
The Haile XV A skull (UF 10909) is in poor condition, and not
enough of the fragments are present to allow complete reconstruction.


FIGURE 15. Nannippus phlegon, left upper molar, occlusal view, UF 17484, Haile
XV A; X 6.15.



Anteroposterior distance
Transverse distance
Crown height
Transverse width, distal end
Anteroposterior width, distal end
Proximal phalanx
Width, proximal end
Width, distal end

UF 17484 UF 17485
17.5 19.6

UNGUAL PHALANX UF 17549a UF 17549b UF 17549c
Total length 43.4 43.6 43.1
Width, distal articular surface 22.7 23.9 22.8

Both left and right molariform tooth rows, however, are complete (Fig.
16) and a portion of the premaxilla has been rebuilt. A small portion
of the occipital region is preserved, but the rest of the skull consists of
isolated fragments.
This skull is referred to the subgenus Equus (Dolichohippus) on the
basis of the well developed parastyles and metastyles, the dilation of the
fossettes, and the more rounded nature of the protocones. Assignment
of extinct North American species, formerly referred to Plesippus, to the
same subgenus as the living Grevy's Zebra follows Skinner and Hibbard
In dental morphology, UF 10909 resembles both E. (D.) shoshonensis
and E. (D.) simplicidens. Gazin (1936) stated that the ranges of mea-
surements of these two species may overlap, although E. (D.) simplici-
dens tends to be smaller. Cheek teeth from Haile XV A (Table 15)
are smaller than any measured by Gazin (1936). They are close to the
size of a specimen of E. (D.) simplicidens described by Hibbard (1941)
from the Blancan Rexroad fauna of Kansas, and they fall at the lower
end of the range given by Howe (1970) for a large sample from the
Broadwater local fauna of Nebraska. The postcranial material is also
small (Table 16), and falls within the range of measurements given by
Gazin (1936) for a specimen of simplicidens.

Equus (Hemionus) cf. calobatus TROXELL

MATERIAL.-UF 17750: upper cheek tooth; UF 17483: lower molar; UF 17482:
M,; UF 17551: left humerus; UF 17552: astragalus; UF 17553: proximal phalanx;
UF 17554: medial phalanx; UF 17555: ungual phalanx.

Vol. 20, No. 3


FIGURE 16. Equus (Dolichohippus) simplicidens, upper dentition, occlusal view,
anterior end upwards, UF 10909, Haile XV A; X 0.80.


chohippus) simplicidens FROM HAILE XV A.1

Left Right
Tooth Length Width Length Width
Tooth row2 158 155
pI 12.2 6.4
P2 32.9 24.7 32.8 24.6
pa 24.1 26.2 24.0 25.2
P4 23.3 24.8 21.9 24.3
M1 22.3 23.8 23.8 25.5
M2 22.0 22.6 22.4 23.9
M3 28.8 23.0 33.2 35.3
SMeasurements taken at grinding surface; width measurements do not include cement.
2 Not including P'.

TABLE 16.-MEASUREMENTS (IN MM) OF Equus (Dolichohippus) simplicidens POST-


Transverse width, distal condyle
Anteroposterior width, distal end
Total length
Anteroposterior width, proximal end
Transverse width, proximal end
Anteroposterior width, distal condyle
Transverse width, distal condyle
Total length
Transverse width, proximal end
Anteroposterior width, proximal end
Transverse width, distal condyle
Anteroposterior width, distal end
Anteroposterior width, proximal end
Transverse width, proximal end
Total length
Total length
Lateral width
Total length
Width, proximal articular surface
Width, distal end
Total length
Width, proximal articular surface
Width, distal end
Total length
Width, proximal articular surface

3 70.1
3 80.7



2 41.2
2 48.5

3 107.7

2 58.0
2 58.8

3 78.5
2 43.7
4 36.4

6 45.5
6 40.7
6 40.9

6 54.4
6 38.5

69.7- 70.5
77.0- 84.0

84.6- 89.7

39.6- 42.8
48.2- 48.8


57.0- 58.9
57.7- 59.8

76.5- 81.2
42.8- 44.5
33.7- 38.5

44.1- 47.4
39.1- 43.2
38.8- 42.8

50.6- 58.8
34.1- 43.3

Vol. 20, No. 3


The three teeth listed above are morphologically different from those
of Equus (Dolichohippus). The fossettes of the upper tooth are much
more complicated, and the lower teeth are quite distinct from those of
E. (Dolichohippus) figured by Gazin (1936), Hibbard (1941), McGrew
(1944), and Skinner and Hibbard (1972). In all the lower molars fig-
ured by the above authors, the median valley divides the isthmus, as
pointed out by Skinner and Hibbard (1972); in the two teeth dis-
cussed here, the median valley approaches the isthmus but does not
divide it. In this respect, the Haile XV A specimens resemble Equus
fraternus, a later Pleistocene ass.
Several postcranial elements in the Haile XV A fauna are probably
also referable to E. (Hemionus) calobatus. They are much smaller than
specimens attributed to E. (D.) simplicidens. Troxell's type and re-
ferred material from the Arkalon (Hibbard 1953) and the Cragin Quarry
local faunas (Schultz 1969) form the bases for comparison. Hibbard
(1956) also reported the medial phalanx of an ass in the Meade Gravel
Member in Meade County, Kansas, with which the Haile XV A medial
phalanx agrees very closely (Table 17).

Tapirus sp.

MATERIAL.-UF 17468: partial crown of a cheek tooth.

Since only one genus of Tapir is known from post-Hemphillian de-
posits of North America, it seems reasonable to ascribe this specimen to
Tapirus. The material serves only to record the presence of this genus
in the fauna. This genus is also recorded in Santa Fe I, another Blancan
locality in Florida (Webb 1974a).

Mylohyus floridanus KINSEY 1974

HOLOTYPE MATERIAL.-UF 18002: left and right mandibular rami and upper left

This species was recently described by Kinsey (1974), based upon
the material from Haile XV A. It represents the oldest (and only
Blancan) record for this genus. The material is refigured (Figs. 17, 18)
and the diagnosis is included here in order to complete the study of the
fauna. Measurements of the holotype are presented in Table 18.


TABLE 17.-MEASUREMENTS (IN MM) OF Equus (Hemionus) cf. calobatus CRANIAL

Anteroposterior distance 25.4
Transverse distance 23.4
M, UF 17482a UF 17482b
Anteroposterior distance 28.7
Transverse distance 11.2 10.7
Total length 255.0
Transverse width, anterior end 88.01
Lateral width, distal condyles 68.1
Anteroposterior width, distal condyles 46.9
Total length 50.4
Greatest width 50.0
Total length 60.6
Width, proximal end 34.8
Width, distal end 30.5
Total length 40.0
Width, proximal articular surface 38.1
Width, distal end 34.8
Total length 45.7
Width, proximal articular surface 33.1

DIAGNOSIS.-Close to Mylohyus nasutus in size and elongation of the
snout. Larger and more robust than Mylohyus fossilis and lacking the
extreme constriction of the symphysis that is diagnostic of that species
(Lundelius 1960). Pre- and post-canine diastema longer than in other
species of Mylohyus. Symphysis much more slender than in Prosthenops.
Coronoid process triangular with straight (not convex) anterior margin.
Total length of the P2-M3 series less than in other Mylohyus, both
absolutely and relative to the overall length of the jaw. M1 and M2
relatively shorter than in other Mylohyus, except M. exoticus (USNM
8876). Anteroposterior diameter of C greater than in all other Mylohyus,
both absolutely and relative to the cheek teeth series. Width of C
greater relative to width of P2 and M1 than in other Mylohyus (Kinsey
DIscussIoN.-Interestingly, Mylohyus is exceedingly rare in most Pleis-
tocene faunas of Florida. For example, a large Late Blancan collection
from Santa Fe I contains only one specimen of this genus, whereas the

Vol. 20, No. 3




It tE''


FIGURE 17. Mylohyus floridanus type, right mandibular ramus, lateral view, UF
18002, Haile XV A; X 0.95.

FIGURE 18. Mylohyus floridanus type, mandibular symphysis, dorsal view, UF
18002, Haile XV A; X 1.0.
other peccary, Platygonus, is very abundant. Similarly, in the Coleman
IIA fauna of Irvingtonian age, Platygonus outnumbers Mylohyus eleven
to one. The presence of Mylohyus at Haile XV A suggests an ecological
distinction, as will be discussed further in the paleoecology section (be-



Left Right

Length, upper canine 15.1
Width, upper canine 11.1
Length, lower canine 14.0
Width, lower canine 10.6
Length, P2 8.5
Anterior width, P2 4.9
Posterior Width, P2 5.1
Length, P, 12.0
Anterior width, P, 9.0
Posterior width, P3 9.1
Length, P4 13.0
Anterior width, P4 10.6
Posterior width, P4 12.6
Length, M, 13.2 13.5
Anterior width, M1 12.6 12.8
Posterior width, M1 12.8 12.7
Length, M2 15.2 15.5
Anterior width, M2 14.0 14.1
Posterior width, M2 13.9 14.1
Length, M3 21.1 21.0
Anterior width, M3 13.1 13.1
Posterior width, M3 11.8 11.9
Length, premolars 33.1
Length, molars 49.9 50.3
Length, molar-premolar series 83.0
Post-canine diastema 80.0
Pre-canine diastema 14.0 13.0
Depth of jaw at M1 41.2 41.0
Thickness of jaw at M1 21.2 21.1
Width, jaw at canines 41.2
Width between canine alveoli 17.6
Least width, symphysis 26.4
Length, articular condyles to anterior end of symphysis 273.0

Measurements after Kinsey (1974).

Hemiauchenia cf. macrocephala (COPE 1893)

MATERIAL.-UF 10900: partial skull; UF 10894: 2 cervical vertebrae; UF 10899:
phalanges (3 medial, 1 ungual).

The Haile XV A skull (UF 10900) (from a juvenile individual) con-
sists of the palate with upper dentition (Fig. 19) and part of the cranium.
The cranial fragment includes parts of the parietal, left squamosal, left
exoccipital, and supraoccipital. This portion of the skull, particularly
the external auditory meatus and the subsquamosal foramina, agrees
very closely with the McKittrick material of this species figured by Stock


FIGURE 19. Hemiauchenia cf. macrocephala, upper dentition, anterior end upwards,
UF 10900, Haile XV A; X 0.93.
(1928). The left DP2 is present, as are the left and right DP3 and DP4;
left and right P3 and P4 are present but unerupted. The complete right
molar series is present, while the left is represented only by the M1.
The Haile XV A specimen is a small llama referable to the genus
formerly called Tanupolama, which is now synonymized with Hemi-
auchenia (Webb 1974b). H. macrocephala is the smallest of the North
American species of Hemiauchenia and has a relatively wider P. than
H. blancoensis. The Haile XV A specimen lacks the critical P4, but the

Vol. 20, No. 3


cf macrocephala FROM HAILE XV A.
Left Right
Length, DP2 10.8
Width, DP2 5.8
Length, DP3 19.7 19.2
Anterior width, DP3 11.51 12.4
Posterior width, DP3 17.3
Length, DP4 19.6 18.8
Anterior width, DP4 17.7 17.8
Posterior width, DP4 19.2 19.1
Length, P3 16.52
Width, P3 9.9
Length, P4 18.82
Width, P4 15.2
Length, M1 26.2 25.7
Anterior width, M1 20.2 21.2
Posterior width, M1 19.8 20.4
Length, M2 22.7
Anterior width, M2 21.2
Posterior width, M2 19.7
Length, M3 26.6
Anterior width, M3 14.2
Posterior width, M3 19.1
Length of deciduous premolar series 50.1
Length of molar series 75.0
Length of tooth row functional at time of demise 1251
Length of permanent tooth row 1101
Palatal width at DP3 34.5
Palatal width at DP4 37.5
Palatal width at M1 47.5
' Estimated.
2 nerupted.

specimen is quite small (Table 19) and tentative species allocation is
based on size.
The occurrence of H. macrocephala in the Haile XV A fauna marks
its earliest record anywhere. A long continuous history for this species
in Florida is indicated by its presence in the early Irvingtonian Inglis IA
fauna (Webb 1974b).
Hemiauchenia was widespread in the Blancan and is known in most
North American faunas of this age, as well as in the Chapadmalalan of
Argentina (Kraglievich 1946).

Odocoileus virginianus ZIMMERMAN

MATERIAL.-UF 10885: 6. partial upper dentitions; UF 17481: right mandible
with P2, DP4, M,, M,; UF 17478: left P4; UF 17477: right M3; UF 10883: 19


Haile XV A Santa Fe I Santa Fe 4A

Lateral width, distal condyles 3 31.0 30.6-31.4 2 28.4 26.2-30.6 1 28.4
Lateral width, proximal end 3 23.5 22.7-24.8 2 27.4 26.6-28.2 1 22.2
Anteroposterior width, proximal end 3 16.7 16.4-17.1 2 20.3 20.2-20.5 1 18.1
Lateral width, proximal end 1 23.2 2 22.4 21.5-23.3 1 20.8 Z
Anteroposterior width, proximal end 1 23.7 2 24.8 22.9-26.6 1 23.2
Length 7 41.0 37.7-43.2 10 43.5 39.4-48.9
Width, proximal end 7 13.4 12.2-13.9 10 13.3 11.6-15.2

Inglis IA

Lateral width, distal condyles
Lateral width, proximal end
Anteroposterior width, proximal end
Lateral width, proximal end
Anteroposterior width, proximal end
Width, proximal end

4 32.9 31.3-34.4

1 29.6
1 20.7

3 25.2 24.6-25.5
3 26.5 26.0-27.4

1 46.6
1 15.4

Coleman IIA

7 30.4 28.2-33.2

1 24.3
1 23.8

7 23.6 22.5-24.8
7 25.2 23.6-35.8

7 43.4 40.3-44.4
7 13.0 12.2-13.8

Reddick I

3 29.6 28.6-31.2

2 27.0 27.6-26.4
2 19.5 19.1-20.0

1 23.1 o
1 21.9

4 42.5 40.6-44.4 2
4 14.7 14.3-15.5 9


Devil's Den

Itchetucknee River

Lateral width, distal condyles 8 35.0 33.5-36.7 7 31.4 28.6-34.0
Lateral width, proximal end 12 28.1 27.0-30.2 11 27.2 23.7-29.6
Anteroposterior width, proximal end 12 20.3 18.5-21.3 11 18.4 17.1-21.5
Lateral width, proximal end 11 25.5 24.3-26.9 14 25.6 21.0-26.7
Anteroposterior width, proximal end 11 26.9 25.7-27.8 14 25.7 22.5-29.6
Length 14 41.5 26.1-45.8
Width, proximal end 14 12.7 11.5-13.8


cervical vertebrae; UF 10891: 3 left humeri; UF 10893: right humerus; UF 17479:
3 metacarpals (2 left, 1 right); UF 10887: metatarsal; UF 17480: left astragalus;
UF 10886: 3 podials; UF 10888: 15 phalanges.

The genus Odocoileus is recognized in many of the Blancan faunas
of North America. In most cases, however, the material has been scarce
and species allocation has not been possible.
The definitions of closely related species (and genera) of deer are
very similar, and the taxonomic problem is compounded by the large
amount of individual variation in these forms. The nature of the antlers
and the fourth lower premolars are considered important in cervid tax-
onomy at the generic and subgeneric levels; unfortunately, these features
are also extremely variable. In Simpson's (1928) discussion of Blasto-
cerus extraneus, he pointed out that specimens of Odocoileus studied
approached Blastocerus in character, and some Recent Blastocerus ma-
terial diverged from his specimen toward Odocoileus. The subgenuus
Procoileus (Frick 1937) is based primarily on the characters of the P,.
Examination of a large sample of Blancan Odocoileus teeth from the
Santa Fe I fauna showed individual variation that approached the char-
acters of Blastocerus, Procoileus, and even Craniocerus. Similar, though
less pronounced, examples of individual variation may be observed in
samples of Odocoileus teeth from several later Pleistocene and Recent
sites in Florida. It appears that tooth morphology by itself is too variable
to permit taxonomic assignments at the species level.
Size also has been a criterion for species determination of fossil deer.
Although the Haile XV A deer are relatively small, this is not considered
taxonomically significant, inasmuch as a size comparison of postcranial
elements from later stages of the Pleistocene (Table 20) shows wide size
variation that encompasses the Haile XV A samples. Harlow and Jones
(1965) also showed that in Florida the size of Recent deer fluctuates
significantly from habitat to habitat.
An element by element comparison of the Haile XV A deer material
with specimens of 0. virginianus from other Florida Pleistocene sites,
and with Recent specimens, shows no consistent differences. Conse-
quently, it is considered safest to refer the Haile XV A cervid to Odocoil-
eus virginianus.

North American vertebrate paleontologists, including Schultz (1938),
Wilson (1938), and McGrew (1944), have long recognized the Blancan
age on the basis of the presence and absence of certain mammalian
genera. McGrew (1944) summarized the criteria as follows:

Vol. 20, No. 3


1) the absence of typically Pliocene genera;
2) the presence of characteristically Middle and Late Pleistocene genera;
3) the presence of certain genera that survived from the Hemphillian but
which did not live past the Blancan;
4) the presence of certain genera that were limited to the Blancan; and
5) the absence of certain genera that did not make their appearance until after
the close of the Blancan.

Each of the above criteria may be examined with regard to the Haile
XV A fauna:
1) There are no typically Hemphillian genera represented in Haile XV A.
Kraglievichia may be of Pliocene age in South America, but it is Blancan
or later in North America. It should be pointed out here that such negative
evidence is never conclusive and is particularly dangerous with regard to
the Haile XV A fauna. The small number of taxa found probably indicates
that the specimens collected represent only a small part of the overall fauna.
2) Several genera that range into the Middle and Late Pleistocene appear in
this fauna; such taxa include Odocoileus, Mylohyus, Smilodon, and Sig-
3) Of the genera that survived the Hemphillian but not the Blancan only
Nannippus is definitely present. The presence in the fauna of a gom-
photheriid may also fit this category, despite the fact that the specimen
cannot be definitely assigned to a genus.
4) Only one subgenus, Dolichohippus (genus Equus), in Haile XV A is re-
stricted to the Blancan in North America. The species Sigmodon medius,
Nannippus phlegon, and Equus (Dolichohippus) simplicidens are wholly
restricted to the Blancan and are widespread through most of North
5) The genera Dasypus and Mylohyus, which previously were thought to have
appeared after the close of the Blancan, are present in Haile XV A. In
view of the peculiar ecological and geographic features of the Haile XV A
fauna, these age extensions are not unexpected and do not affect the hy-
pothesis of an older age suggested by the weight of other evidence. In each
case the species is more primitive than those occurring in younger faunas.

The weight of the mammalian faunal evidence (see Table 21), par-
ticularly the presence of Nannippus, Equus (Dolichohippus), and Sig-
modon medius, points conclusively to a Blancan age. A more refined
correlation of the Haile XV A fauna with other North American Blancan
faunas (i.e. outside of Florida) is not feasible at present because of the
geographic distances and ecological differences. Most Blancan faunas
are located in western North America, but none, other than those in
Florida, are known from the Gulf Coastal Plain. Consequently, the lack
of a more detailed faunal comparison need not imply a great difference
in time. The absence of Kraglievichia and Dasypus from Blancan sites
elsewhere in North America more likely results from geographic rather
than temporal differences. Conversely, a closer similarity to other Blan-
can faunas in Florida (e.g. Santa Fe I) does not indicate a close temporal
correlation. Although definite similarities exist between the Haile XV A
and Santa Fe I faunas, enough differences are present to indicate a differ-


Equus (Dolichohippus)


x C.


X g

? X X



X X X X X X ? X X X




X X X X X X X X X ?

X X XX z
x x x x

Haile XV A

E m E
44 44 -4
1 i ~1 c

1 'o 0v
0: 0 0 0 I
Sg 2 9M 3
4" 44 '4 0 D
E4 E g 9 0
.I s I -. : ~ 4 Im I' I 'I
z c
0 0 .4 z 0 0 5 0 *

I I 0 I0 W Is i I 0 i
0 00 44 u. w m 4 '4u Z4
~~ 0 '"o >d Z F .
g0.M H "s -; --ba'
44 8 0 0 2, ,, E- ~.3 2 '- '
urnjm m OH rnuou~ r n~z


ence in time and probably ecology as well. The Santa Fe I camel be-
longs to the typically Blancan species Hemiauchenia blancoensis, whereas
the Haile XV A form most closely resembles H. macrocephala. This
probably indicates a younger age for Haile XV A, as H. macrocephala
is characteristic of the Irvingtonian and Rancholabrean ages elsewhere.
Certain taxa found in the Santa Fe I fauna that might be expected
at Haile XV A do not occur there. One of the most notable absences
is the typically Blancan hyaenoid dog, Borophagus, although in all like-
lihood this represents a sampling error. The absence of Castoroides and
Hydrochoerus from Haile XV A probably can be explained in the same
manner. Other differences between the faunas of Haile XV A and Santa
Fe I are most likely ecologically induced, as will be discussed in the
next section.
Among the best represented animals in the Haile XV A fauna are the
edentates, a group of undoubted South American origin. Of the three
genera at Haile XV A, only Glossotherium has been recognized pre-
viously in North American Blancan faunas, being present in the Blancan
faunas of Texas and Nebraska (Table 21). Dasypus is known only
from later deposits elsewhere in North America, and Kraglievichia has
not heretofore been reported from this continent, although its probable
descendant, Pampatherium (=Holmesina), has been found in Irving-
tonian and Rancholabrean deposits.
This early record of a major influx of edentates from South America
is further indication of the late Blancan age of Haile XV A. The small
chlamythere from Haile XV A shows a remarkable resemblance to the
species Kraglievichia paranensis from Late Pliocene deposits of South
America. Although no authentic Chapadmalalan (Early Pleistocene)
species is known, it probably lived in regions more tropical than Argen-
tina and probably also was physically similar to the Florida material.
Florida and Argentina apparently represent the northern and southern
limits of the range for this form, and it seems safe to conclude that con-
temporaneous populations occupying the middle portion of the range
were broadly similar to those at the extremes. The close resemblance
of the Haile XV A Glossotherium to that from the Chapadmalalan stage
also supports this correlation.
This late Blancan influx of edentates into Florida adds a new dimen-
sion to the previous correlation between the Blancan stage of North
American and the Chapadmalalan stage of South America. This correla-
tion had previously been based upon the appearance of a few Nearctic
species in the Chapadmalalan and Uquian of South America (Patterson
and Pascual 1968, Simpson 1969). Haile XV A adds the obverse re-
lationships. North America received an important contingent of Neo-


tropical taxa in the Blancan. The only Nearctic taxa in the Haile XV A
fauna that closely resemble species in the Chapadmalal fauna are Smilo-
don and Hemiauchenia. The combination of correlative taxa of both
Nearctic and Neotropical origin makes the resemblance between the
Blancan fauna of Florida and the Chapadmalalan fauna of Argentina
particularly strong.

In addition to faunal correlation, the effects of sea level changes
caused by Pleistocene glaciation may be employed as an aid in dating
Florida Pleistocene deposits. Glacial eustatic fluctuations of sea level
have resulted in "terraces" or "abandoned shore lines," which are domi-
nant geomorphic features of the Florida landscape (Cooke 1945, Mac-
Neill 1949, White 1958, Alt and Brooks 1965). The location of a deposit
that can be correlated with these features can be quite helpful in age
determination. For example, a terrestrial deposit that is presently lying
at or below sea level must have been laid down at a time when sea level
was as low or lower than it is now. This would be an indication that the
deposit was laid down during glacial times, as these were the times of
low sea level. Once this is determined, the faunal elements may indi-
cate which glacial stage is represented.
Inland deposits at higher elevations present a problem in that they
could have been deposited either during a glacial or an interglacial stage.
If it can be shown that a particular inland deposit was coastal at its
time of deposition, then it must represent an interglacial stage, when sea
level was high. Haile XV A stands at an elevation of 90 ft above sea
level. It contains shark vertebrae that are similar in preservation to the
other vertebrate fossils in the site (and therefore are probably not in-
trusive). This indicates the close proximity of the sea during the time of
deposition. Remains of other fish species also suggest at least estuarine
conditions. Haile XV A could represent either an interglacial stage or a
preglacial wet interval during the Blancan; a preglacial age seems prob-
able on biostratigraphic grounds.

Among the lower vertebrate remains from Haile XV A are numerous
lamnoid type shark vertebrae. Their preservation does not differ from
that of other vertebrate hard parts in the site, and thus they cannot be
interpreted as intrusive from adjacent Eocene limestones. (Sharks are
known in the Ocala Formation, but not abundantly and mainly from
teeth.) A variety of bony fishes of both freshwater and marine taxa
await further study. These include Amia, Lepisosteus, Centropomus,

Vol. 20, No. 3


Bagre, Ictalurus, and numerous other as yet unidentified species. Am-
phibians are represented by both anurans and urodeles. The turtles from
Haile XV A include the aquatic forms Chrysemys platymarginata, Tri-
onyx, Kinosternon, and Chelydra, and the terrestrial box turtle Terrapene
carolina. The material of Chrysemys platymarginata (Weaver and Rob-
ertson 1967) includes numerous complete shells. They all occur in up-
right position, but facing in various directions, and are most abundant
in the uppermost green clay unit. Most of the birds at Haile XV A are
aquatic and include a duck, three herons, and a grebe (Campbell 1976).
Although the lower vertebrate fauna has not been studied in detail, this
preliminary analysis is enough to indicate an aquatic environment near
the sea.
The mammalian taxa may be separated for the most part into two
ecological components, a smaller group of aquatic species and a larger
group of terrestrial species. Castor inhabits medium to larger permanent
bodies of flowing water. Pteronura, the probable descendant of Sather-
ium, inhabits streams from the Guianas to Argentina (Walker 1975). Its
surrounding terrestrial habitat is probably secondary to its stream habi-
tat, as it lives in suitable rivers that flow through a variety of terrestrial
areas. These mammals are thus ecologically associated with the turtles,
fishes, and water birds as a primary inhabitants of the aquatic site.
The abundance of large terrestrial animals and the relative scarcity
of smaller terrestrial ones strongly suggests a spring-head habitat. The
presence of a few of the denser bones (and especially teeth) of Sig-
modon, Scalopus, Cryptotis, and Cryptopterus must be ascribed to for-
tuitous burial. The only other small mammals in the fauna are Castor
and Satherium, which are normally aquatic in nature and thus may be
expected abundantly in the stream. All the other mammals in the Haile
XV A fauna are large terrestrial forms, which were probably trapped
as they drank by the treacherous slopes and abrupt limestone ledges that
often surround Florida springs.
We may next consider the nature of this terrestrial faunal sample.
The terrestrial animals consist of Terrapene, the box turtle, one species
of quail, one turkey (Campbell 1976), a few small mammals, and a con-
siderable variety of large mammals. The presence of eurytopic large
forms, such as Smilodon, Hemiauchenia, two subgenera of Equus, Nan-
nippus, Odocoileus, and of the small mammals Cryptotis, Scalopus, and
Sylvilagus tell us little about the local paleoecology. These forms are
present in many Blancan local faunas of North America and probably
ranged through many habitats.
Several taxa in the Haile XV A fauna, however, strongly suggest the
presence of a forest community adjacent to the site of deposition. The


new species of Mylohyus lends strong support to this interpretation.
Lundelius (1960) reviewed evidence that indicates a forest habitat for
Mylohyus. He also included Castor, Tapirus, and Glaucomys (among
others) as forest forms. Both Castor and Tapirus are present in the
Haile XV A fauna, and the flying squirrel Cryptopterus (like Glaucomys)
certainly indicates a forest environment. Glossotherium may also have
favored forested or partly forested areas.
Dasypus and Kraglievichia probably were restricted in North America
to the Gulf Coastal Plain by their need for moderate to warm tempera-
tures and high rainfall. This may explain their absence from all other
North American Blancan faunas. In South America, Kraglievichia seems
to have been largely restricted to the northern tropical part of the con-
tinent. These edentates suggest a tropical or subtropical climate in
central Florida during the Latest Pliocene.
In summary both the geological and faunal evidence seem to indicate
that the Haile XV A site lay at the springhead of a coastal stream that
flowed through a tropical or subtropical forest or forest savanna.

The Haile XV A fauna is the first Blancan fauna to be described
from the Gulf Coastal Plain. Its unusual faunal assemblage (compared
to Blancan faunas in western North America) apparently is a reflection
of its location. Few American Blancan faunas are from coastal areas,
and no others are so close to the Middle American tropics.
The most striking distinction of the Haile XV A fauna is the diversity
of edentates. Evidently it was in contact with the tropical corridor
through which South America emigrants moved. Of the edentates,
only Glossotherium was not restricted to the tropical portion of North
America, as indicated by its presence in the Blanco (Texas) and Broad-
water (Nebraska) faunas. This genus probably was also widely dis-
tributed on the South American continent, as the presence of G. chap-
admalensis in Argentina indicates. On the other hand, Kraglievichia
probably was restricted to the tropical or subtropical parts of North
America during Blancan time, as indicated by its absence from all
other North American Blancan faunas. Its presence in earlier Pliocene
deposits of Argentina suggests that its range extended southward into
more temperate climes in South America, but later it seems to have re-
treated tropicad in that continent also. Apparently the later Pleistocene
evolution from Kraglievichia to Pampatherium involved, among other
things, an increased tolerance to colder climate, which perhaps was cor-
related with the greater size of Pampatherium. Pampatherium was
able, in the course of the Pleistocene, to disperse northwestward from

Vol. 20, No. 3


the Gulf Coastal Plain throughout most of temperate North America; in
South America it spread southward throughout Argentina.
The genus Dasypus seems to have been even more closely restricted
to tropical regions than either Glossotherium or Kraglievichia. It also
was able to move out of the Gulf Coastal Plain later in the Pleistocene
(as Dasypus bellus), reaching as far west as Texas and as far north as
West Virginia (Guilday and McCrady 1966), Tennessee (Guilday et al.
1969), and Missouri (Simpson 1949). The probable mechanism permit-
ting its tolerance to colder climate may have been an increase in size,
as was true of Pampatherium. Its drastic size decrease in latest Pleisto-
cene time (assuming D. bellus is closely related to D. novemcinctus)
could account for its present limited southern distribution (Humphrey
1974). Dasypus bellus is not known from South American fossil de-
posits, although there is little doubt that the genus originated on that
continent. This suggests that its range was similarly restricted to tropical
areas on that continent, where fossil sites are rare and little known.
The unusual distributional pattern of Cryptopterus (Latest Pliocene
of Florida and trans-Beringean of West Germany) indicates a wide distri-
bution of large Group I flying squirrels. Presumably this group was re-
stricted in the New World to the eastern deciduous forests of North
America. James (1963) reported a smaller simple-toothed group of
sciuropterines in Miocene deposits of California, but no other large
complex-toothed specimens are known from the New World. Presum-
ably Cryptopterus ranged from the eastern deciduous forests of North
America through similar forests in Alaska and Siberia into Central
Europe. The presence of Pteromys at Choukoutien (Young 1934) adds
another very late Cenozoic record of a large flying squirrel from a temper-
ate region, but it is not closely related to Cryptopterus (Mein 1970).
The close resemblance between the Florida and West German specimens
suggests rapid dispersal to the New World during late Hemphillian or
Blancan time. Obviously this fortuitous record from the Blancan of Flor-
ida is peripheral to the main theatre of sciuropterine evolution in the
Old World tropics.

The Haile XV A mammals are from one of the few Blancan faunas
on the Gulf Coastal Plain. It is the only such fauna that has been ex-
tensively studied to date and also the only one from a subtropical forest
Included among the mammals from this site is Glossotherium chap-
admalensis, previously known only from Argentina. It is possible that
this form gave rise to G. robustus in South America and G. harlani in


North America. Apparently, Glossotherium was able to spread through-
out much of temperate North America, whereas the other edentates
represented in the Haile XV A fauna were restricted to subtropical re-
gions during most of the Pleistocene.
The occurrence of a new species of Kraglievichia at Haile XV A and
Santa Fe I is the first record of this chlamythere genus in North America.
The presence of this species, which is closely related to K. paranensis,
strengthens the correlation between North and South American faunas
of Blancan and Chapadmalalan ages respectively.
The oldest evidence of Dasypus bellus occurs at Haile XV A. This
species or its immediate ancestors undoubtedly originated in South
The giant flying squirrel, Cryptopterus, gives the local fauna an un-
expected link with Old World Tertiary forest faunas. The new species
from Haile XV A was among the last (Late Pliocene) dispersals of Old
World subtropical forest dwellers to North America (Repenning 1965).
The presence of Satherium in the Haile XV A fauna marks the first
Florida record of this large aquatic otter. It adds yet another link to
the neotropical fauna, as it is closely related to Pteronura, the giant
otter of South America.
The presence of Mylohyus floridanus at Haile XV A marks the earliest
record for this genus, although its ancestor, Prosthennops, is known from
the Pliocene of North America. It probably occurred in subtropical
forests, in contrast to the contemporary genus Platygonus.
If the tentative assignment of the Haile XV A camelid to Hemiau-
chenia macrocephala is correct, it marks the earliest record for this species
anywhere. The typical Blancan species, H. blancoensis, occurs at Santa
Fe I, suggesting an age difference between the Florida Blancan localities.
The remaining members of the Haile XV A fauna are fairly typical
representatives of the North American Blancan fauna. Such character-
istic taxa as Equus (Dolichohippus) simplicidens, Nannippus phlegon,
and Sigmodon medius substantiates a Blancan age assignment for Haile
The presence of sharks and brackish-water fishes indicates that the
sea stood near Haile during the time that Haile XV A was deposited.
The presumed sea level, which was 90 ft higher that that of today, is
believed to represent a pre-glacial high sea level.
Although the bulk of the vertebrate fauna at Haile XV A indicates
stream conditions, an adjacent terrestrial community is also well repre-
sented. Of the mammals, only Satherium and Castor represent the
aquatic community. The terrestrial community includes several clear

Vol. 20, No. 3


indicators of a tropical forest situation, such as Mylohyus, Cryptopterus,
three kinds of edentates, and Tapirus. The others are eurytopic small
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